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FIELD-NATURALIST
Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada
Voi. XXXII, No. 5. NOVEMBER, 1918.
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Centennial Year
Volume 93, Number 1 January-March 1979
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
Patrons
Their Excellencies the Governor General and Madame Jules Léger
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural
heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse
information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving,
maintaining or restoring environments of high quality for living things.
The Members of Council are listed on the inside back cover.
The Canadian Field-Naturalist
The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas
expressed in this journal, however, are private and do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or
any other agency.
Editor: Lorraine C. Smith
Assistant to the Editor: Donald A. Smith Book Review Editor: J. Wilson Eedy
Associate Editors
C. D. Bird A. J. Erskine David P. Scott
E. L. Bousfield Charles Jonkel Stephen M. Smith
Francis R. Cook Charles J. Krebs Robert E. Wrigley
George H. La Roi
Copy Editor: Marilyn D. Dadswell Chairman, Publications Committee: J. K. Strang
Production Manager: Pauline A. Smith Business Manager: W. J. Cody
Subscriptions and Membership
Subscription rates for individuals are $10 per calendar year. Libraries and other institutions may subscribe at the rate
of $20 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $10 includes a subscription to The
Canadian Field- Naturalist. Subscriptions, applications for membership, notices of changes of address, and undeliverable
copies should be mailed to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada KIY 4J5.
Second Class Mail Registration No. 0527 — Return Postage Guaranteed.
Back Numbers
Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field- Naturalists’ Club, 1879-
1886, and The Ottawa Naturalist, 1887-1919, may be purchased from the Business Manager.
Business Manager: Mr. W. J. Cody, Box 3264, Postal Station C, Ottawa, Ontario, Canada KIY 4J5
Book Review Editor: Dr. J. Wilson Eedy, R.R. !, Moffat, Ontario LOP 1J0
Coordinator, The Biological Flora of Canada: Dr. George H. La Roi, Forestry Sciences Laboratory, 3200 Jefferson Way,
Corvallis, Oregon, USA 97731 (address valid until August 1979).
Address manuscripts on birds to the Associate Editor for Ornithology:
Dr. A. J. Erskine, Canadian Wildlife Service, Box 1590, Sackville, New Brunswick E0OA 3C0
All other material intended for publication should be addressed to the Editor:
Dr. Lorraine C. Smith, R. R. 3, Stittsville, Ontario, Canada KOA 3G0
Urgent telephone calls may be made to the Editor's office (613-996-5840), the office of the Assistant to the Editor (613-231-
4304), or their home on evenings and weekends (613-836-1460), or to the Business Manager's office (613-995-9461).
Cover: Since the Club was founded in 1879, it has published a journal under three successive names. Covers bearing the two
names preceding the present one are illustrated. Left, Ortawa Field- Naturalists’ Club, Transactions, published for 1879
to 1886. Right, The Ottawa Naturalist, published from 1887 to 1919; it bore this cover design only between April 1918 and
March 1919.
THE CANADIAN
FIELD-NATURALIST
Volume 93
1979
CENTENNIAL YEAR
THE OTTAWA FIELD-NATURALISTS’ CLUB
OTTAWA CANADA
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ae
The Canadian Field-Naturalist
Volume 93, Number | January-March 1979
One Hundred Years in Perspective — the Changing
Roles and Objectives of
The Ottawa Field-Naturalists’ Club
ROGER A. FOXALL
President, The Ottawa Field-Naturalists’ Club, January 1977-January 1979
Following an invitation to members of the Ottawa Literary and Scientific Society, “fully forty
gentlemen attended the meeting held on the 19th March, 1879. After a lengthy discussion as to the
form the organization should take, the Ottawa Field-Naturalists’ Club was born.”
The quotation above is from the first historical sketch of the Club, taken from the 1880 Annual
Report when the Club was fully one year old. Since then there have been several articles describing
the many memorable people and events that add up to the varied and fascinating 100-year history of
our Club. A series of historical articles is currently being published in Trail & Landscape. The aim of
this message is not to provide another summary of people or events, but to review instead the ways in
which the Club’s functional role and objectives have changed over time and to present a personal
view of our present role and objectives in terms of future needs.
In the beginning, the objective of the Club was stated simply as “the study of the Natural History of
this locality.” The raison d’étre, however, was more explicit in an 1884 summary:
“The value of these studies cannot be over-estimated for, from their very nature, they are
most beneficial both to mind and body, while, from an economical point of view they are no
less important . . . . The future of Ottawa is in large measure dependent upon the
development of the vast stores of minerals contained in the surrounding districts, and
which are as yet almost unexplored. The manifold uses and demand for all vegetable
products make it necessary that the nature and habits of plants should be closely studied, as
well as those of the countless myriads of insect foes which deprive the cultivator of so much
of his hard-earned gains... . It gives very great satisfaction to the members of the Council
to find that the Club is being gradually recognized as a source of reliable reference on all
these matters.”
So the objective in part was the study of natural history to assist the economical development of
the Ottawa area. Times have indeed changed! A specific example of success in this direction was the
advice given to the Ottawa Granite Company that within a few miles of the city there existed an
outcrop of quartzite deposits just as good as quartzite then being imported from New York State.
This advice led to successful commercial exploitation of the deposits.
Although economic development of the Ottawa area was one aim of the Club, another was to
popularize the study of the different branches of natural history and thereby increase appreciation of
nature. On the local scene this entailed programs of organized studies, excursions, lectures, and
generally spreading the good word. By the thirtieth anniversary, Club membership had increased
from the initial 40 to over 300. Increasingly, however, the Club’s successes led to recognition
throughout the Dominion, and its role began to broaden from that of a strictly local club towards
2 THE CANADIAN FIELD-NATURALIST Vol. 93
that of a national society. In an 1893 The Ottawa Naturalist, the editor found it necessary to state,
“Although the scope of the Club’s work has been gradually widened to receive the benefits of
investigations made by its members wherever they may be located, the special work for which it was
organized must stili merit the chief attention, and although much has been observed and recorded of
the Natural History of Ottawa, there still remains vastly more to be done.” But the transformation
accelerated! A series of 42 nature study articles was written for The Ottawa Naturalist by leading
Club members, and between 1903 and 1908 a total of 238000 separate copies was distributed to
schools and colleges across Canada. Such was the success and expanding influence of the Club that
Ottawa’s Evening Journal (although perhaps a little biased) found it appropriate in 1908 to state that
“today The Ottawa Field-Naturalists’ Club is the greatest institution of the kind in the entire
continent of North America.”
Partly because of the Club’s campaigns and the travels of Club members, but mainly because of a
general increasing interest in nature, similar organizations were established in several cities across
Canada. Increasingly, the need developed for a publication covering the natural history of all of the
Dominion, until in 1918 the Foreword to the last volume (32) of The Ottawa Naturalist announced:
“The time has come, however, when a local periodical of this nature is inadequate and the
Dominion requires a more creditable and representative publication for the record and
dissemination of the results of scientific research. The Ottawa-Naturalist, with its already
established position, long and honorable history and scientific standing, seems a logical
nucleus from which such a publication should be developed.”
The first issue of The Canadian Field- Naturalist was published in April 1919. This event in the Club’s
40th year marked the end of a lengthy transition and formalized the dual role of the Club — a local
natural history club and a national society. Somewhat earlier, in October 1912, the objectives of the
Club had been expanded significantly to reflect the two roles:
To foster an acquaintance with and a love for nature; to study especially the natural history
of the Ottawa District; to encourage investigation and to publish the results of original
research in all departments of natural history; to arrange for out-of-door excursions during
the summer months; to provide free lecture courses during the winter months; and ina
general way to render assistance to students or others interested in Nature Study.
With one significant exception, to which I will refer below, these objectives remained unchanged
until 1972.
Unfortunately the success of the Club’s first 40 years was to be followed by a 30-year period during
which many difficulties were encountered. Until about 1923, the Club received a grant from the
Ontario Department of Education that contributed significantly to the sustenance of The Ottawa
Naturalist and later The Canadian Field-Naturalist. \ronically, post-war economies led to
cancellation of the grant when it was needed most, for the creation of The Canadian Field- Naturalist
placed a significant burden of responsibility on the Club and all its members. Without the grant,
many more members were needed to provide financial stability. But the growth was too slow. In 1934
P. A. Taverner, Chairman of a Special Committee on Membership, issued an appeal to all Canadian
naturalists to boost membership. A reduction in size, from 24 to 16 pages an issue, and other
parsimonious economies had been tried, but still the year-end balance was negative. The journal
survived, of course, but for many years the fight for The Canadian Field-Naturalist’s survival
represented the principal objective of The Ottawa Field-Naturalists’ Club.
The difficulties in maintaining the national role were paralleled by difficulties with the local role.
Here, analysis is more difficult. We can perhaps rationalize the decreasing enthusiasm in terms of
two gradual developments. The early years of systematic, detailed studies of local natural history
must have represented a frontier-like challenge — new knowledge was easily acquired and each
discovery, be it a new species of plant, insect, bird, or butterfly, contributed to the developing picture.
But the acquisition of new knowledge makes further discoveries just that much more difficult.
Members would have to travel further and look harder, and greater expertise was needed. But why
1979 FOXALL: PRESIDENT’S MESSAGE 3
try so hard? The initial objective, to assist in the economic development of Ottawa, was surely less
important now. The urgency of acquiring knowledge in support of conservation and preservation to
slow down development was not yet recognized. For whatever reasons, it is a fact that emphasis on
studying the local natural history declined to the point that in 1947, the objective “to study especially
the natural history of the Ottawa District” was removed from the Constitution. Although excursions
and lecture programs continued to foster and sustain members’ interests in local natural history, the
membership grew slowly and even declined for several years.
In retrospect it seems likely that the Club’s surviving the difficult 30 years can be attributed to its
dual role. For without the local membership, The Canadian Field-Naturalist might have failed
financially, and without the responsibility of continuing The Canadian Field-Naturalist, the local
club might have failed for lack of sufficient motivation.
The last 30 years, 1949-1979, have also been characterized by significant changes, but the story is a
happier one. The Canadian Field- Naturalist has steadily developed into a scientific journal of both
national and international repute — a journal almost unique in the world for its breadth and
standard of scientific natural history reporting. The content has evolved also. The studies reported
have progressively become more sophisticated as new knowledge requires increasingly more detailed
examination of natural phenomena. Other, less formal journals and club magazines are now
published and these provide media for reporting studies and findings of more local importance.
There is no doubt that these evolutions have resulted in The Canadian Field- Naturalist becoming of
lesser interest to many local members. But the need for The Canadian Field-Naturalist is at
least as great today as it was in 1919, and, through its continuing support, The Ottawa Field-
Naturalist’s Club is playing a very valuable role in the reporting of Canadian natural history.
The continuing decrease of Ottawan content in The Canadian Field- Naturalist and the need to
regenerate the local Club, led to the start, in 1949, of a Club Newsletter for local members. Steadily
the pendulum swung back as successive Presidents and Councils worked very hard to rebuild. It was
not easy, as illustrated by the anguished appeals in the Newsletter of Chairmen of Excursions and
Lectures urging members to attend the many walks and talks that were being arranged. But the
rebuilding succeeded and by the late 1960s the local Club had regained much of the status and drive
of the early period. As a Canadian Centennial project, the Club upgraded the Newsletter to become
the magazine Trail & Landscape. For the last eleven years, the staff of Trail & Landscape has
succeeded in producing an excellent, informative magazine that focusses on local sightings, events,
and issues. Sometimes serious, sometimes lighthearted, but always highly readable, Trail &
Landscape plays an important role in generating and maintaining the enthusiasm of local members.
Somewhat surprisingly, in retrospect, natural history organizations throughout North America
were slow to recognize that man’s rapid development and exploitation represented a serious threat to
all natural environments and living things. In the beginning, The Ottawa Field-Naturalists’ Club had ~
worked in support of economic development. When, later, favorite areas such as Dow’s Swamp were
taken from us, no organized howls of protest were heard. Such things were taken for granted as a
normal result of development. But by the mid-1960s organizations everywhere recognized the threat
to what had seemed before to be an apparently infinite environment. The battle began to preserve for
future generations those areas that had so far survived exploitation, to control thoughtless pollution,
and to prevent the indiscriminate use of chemicals that threatened wildlife. In 1972, the Club’s
objectives were rewritten to reflect the new motivation:
To promote the appreciation, preservation and conservation of Canada’s natural
heritage; to encourage investigation and publish the results of research in all fields of
natural history and to diffuse information on these fields as widely as possible:
to support and cooperate with organizations engaged in preserving, maintaining or
restoring environments of high quality for living things.
On the national scene, the new emphasis is being supported by the publication in The Canadian
Field- Naturalist of, for example, excellent papers on the effects of chemicals such as PCBs in the
environment, and the internationally acclaimed issues on the status of the Peregrine Falcon. The
4 THE CANADIAN FIELD-NATURALIST Vol. 93
local Club established a Conservation Committee of Council and became heavily involved in the
battles to defend the Mer Bleue and Gatineau Park, and more recently to defend many other natural
areas of the Ottawa-Carleton Region.
Today, the Club continues its dual role, publishing The Canadian Field- Naturalist on behalf of all
Canadian naturalists, and performing the functions of a typical local natural history club —
arranging excursions and lectures, publishing a local magazine, and attempting strenuously to
protect from further development many areas of significance to natural history.
A successful one hundred years? Certainly! The Club’s founders would surely be pleased with what
transpired from that first meeting on 19 March 1879. The Club has evolved to meet the needs of
naturalists over a period of immense change in Canada, thousands of people have benefited directly
from their association with it, and many more thousands have benefited indirectly from its
achievements. More could have been done, particularly on the local front, for it is surely ironic that a
club formed one hundred years ago to study Ottawa’s natural history has been forced to scramble
hurriedly during the last few years trying to acquire sufficient knowledge of local areas (some only
ten miles from Parliament Hill) to support their preservation and conversation.
Although the celebration of our Centennial Year will be largely retrospective, some time should be
spent thinking about and discussing our current roles and objectives in terms of future needs. Some
predictions can be made with a reasonable degree of confidence. These are:
— the need for The Canadian Field- Naturalist, or an equivalent journal, will continue into
the foreseeable future:
— the pressure on natural environments and living things will continue to increase, in the
short term at least;
— the potential for increasing public interest in and concern for natural history will
increase as the amount of leisure time continues to increase: and
— in the longer term, leisure activities, if uncontrolled, will become a serious threat to
natural environments.
Let us now examine our objectives in these terms. The second objective, “to encourage
investigation and publish the results of research in all fields of natural history and to diffuse
information on these fields as widely as possible,” is certainly appropriate. Although several local
Club members have expressed the opinion that The Canadian Field- Naturalist has become too
“professional” or too “scientific” to be published by a club largely made up of amateur naturalists, I
remain convinced that the Club should continue to publish The Canadian Field-Naturalist on
behalf of all Canadian naturalists at least until it is demonstrated that another organization could
provide it a better home.
Similarly, the third objective, “to support and cooperate with organizations engaged in preserving,
maintaining or restoring environments of high quality for living things,” will continue to be
appropriate. Our recent performance in support of this objective has not been outstanding and the
Club should concentrate on becoming more involved with other organizations. To attempt to do this
effectively on a national scale would be unrealistic, but more could and should be done to promote
effective cooperation with organizations in Ontario and Quebec.
The first objective, “To promote the appreciation, preservation and conservation of Canada’s
natural heritage,” sounds impressive but is as a separate objective, the least realistic for The Ottawa
Field-Naturalists’ Club today or in the future. We do support it to some extent by publishing The
Canadian Field-Naturalist and by cooperating with other organizations, but some modification of
the objective appears necessary if the Club is not to pretend it is trying to duplicate the function of a
large national conservation organization such as the Canadian Nature Federation. This is not to say
that I feel The Ottawa Field-Naturalists’ Club should become overly parochial, but I do think that
the stated objectives of any organization must represent specific, realistic challenges that can be
pursued effectively.
Finally, | would urge the Club to reintroduce the objective “to study especially the natural history
of the Ottawa District,” with this or some similar wording, to reflect the localized nature of most of
1979 FOXALL: PRESIDENT’S MESSAGE 5
our activities and the need to continue increasing our knowledge of the district. During the
conservation battles of the last few years, local planning bodies have come to recognize the Club as
the authoritative source for much of the data on which to base decisions affecting the natural
environment. This recognition is due entirely to the hard work of relatively few members. Past
appeals to members to help in this endeavor by turning out for inventory-type excursions have largely
failed. If the Club is to succeed in meeting the challenges for the future based on the predictions listed
above, it must surely focus again on the original objective, chosen on 19 March 1879 — “the study of
the Natural History of this locality.”
Biology — The Unknown Science?
YORKE EDWARDS
British Columbia Provincial Museum, Victoria, British Columbia V8V 1X4
Biology is a science largely unknown to the people of the world. If life is the most important
“thing” on Earth, this is a strange condition. The disturbing result is that not only is the man in the
street largely ignorant in all things biological, but also that decisions on how man will treat our life-
support system — Planet Earth — are made without biological understandings. Biology is largely
missing as general knowledge, hence also as an automatic aid to intelligent decisions. When, on
occasion, biology does happen to be given its day, there is no background of accepted status to give it
weight. Biology has little hope of being valued or even understood if it is to surface in public affairs
only at moments of crisis.
Scientists do a remarkably good job of keeping the public uninformed. This is partly because the
public is rather narrow-minded about what is important, which creates some reluctance on the part
of scientists to become too visible. But then, of course, a more informed public might be more
understanding. When we add to this picture the high proportion of scientists unable to be very
informative about their work to almost everyone, the reasons for science’s isolation from the man in
the street are clear and appear to be formidable.
The poor record of scientists being intelligible and convincing in public seems to be programmed
that way by the nature of both today’s science and today’s scientists. Science today is largely
concerned with exploring smaller and smaller bits of our world, with things beyond the awareness of
everyday human experience. Scientists lead this trend, but few are helping to put the bits together
into more understandable wholes. It is the era of the specialist. By contrast the generalist in science
has become rare, which I am convinced endangers both science and the world.
The generalist in science is a necessary companion to the abundant specialist who is traditionally
learning “more and more about less and less.” The need is not just to assemble what we know about
the world into forms meaningful to many, it is also a matter of possessing the resulting wisdom
necessary for survival.
Man’s wildly compounding knowledge has him more and more in control of his support system
Earth in an accelerating use of this control that is quite out of control. Science feeds the largely
destructive trend. Few people, including most scientists, can glimpse what is really happening, for the
quality and quantity of the effects are beyond the sensory capacities of most people. Understanding,
acquired from those few who do understand, is the only hope of self control, understanding that
comes from broad insights into the world and how it functions. Science must do a much better job of
arranging “more and more of less and less” so that bit plus bit becomes a somewhat visible, somewhat
accurate, somewhat understandable whole.
Many scientists consider the discovery of truth to be their sole justification. For example, they
regard the basic discoveries in atomic physics as standing alone; atomic warheads are therefore other
men’s burdens, not those of the pure scientists. This assumes it not true that achieving atomic fission
made Hiroshima’s devastation probable, even inevitable in the political climate of the time.
In most respects all of us have minds not much changed from those of ancestors in Ice Age caves.
The products of science available to us all are power in the hands of ignorance. If science put the
power there, and continues to do so without adding very effectively the controls of understanding, we
are surely building systems for our own destruction. Only science can correct this situation. We need
a new morality in the sciences, which would result in major efforts to make science and its influence
on Earth understandable to those outside the ivory towers, and to those in them too, for most
scientists do not understand most science.
There is a modern myth in our culture that the scientist is among the most brilliant and wise of
men, understanding the world as few others do. Some scientists believe this. But there are many
6
1979 EDWARDS: GUEST EDITORIAL 7
people who do not, and in fact, from his specializations, and from his frequent mental and physical
isolation from the worlds that most people experience, there is an emerging image in our society of
the scientist being rather dangerous.
Inevitably, dedicated specialization results in tunnel vision. Neither the specialization nor the
resulting narrow view of the world are by any means confined to scientists in our society, nor is the
trend for these many specializations to be increasingly specialized. But they are relatively
unimportant outside of science. It is specialization within science itself that can keep people awake at
night.
Science has given us formerly unbelievable powers and capabilities at the price of mushrooming
humanity in a disintegrating support system. If it is not already too late, the end of irresponsible
science is the only cure, and then only if irresponsibility is replaced by responsibility, not just for
tomorrow’s new problems but for yesterday’s legacy of old ones as well.
It is fashionable to say that “It was the side-effects that got us.” There can be no clearer evidence of
our narrow views. “Side-effects” are a tunnel-vision illusion, for the broad view will see only
BCILCCtS:
I have begun this essay with the broadest possible sweeps quite by plan. My resulting
generalizations contain many of the “part truths” and “too general” sorts of “facts” that drive most
scientists into ever deeper specializations with their attractively simple foci of study. In disciplines
properly requiring the greatest possible degrees of accuracy, the uncontrolled drift of scientists
toward the high accuracy areas is perhaps predictable. But I wonder if this is both science’s blind
alley, and Earth’s sentence to increasing impoverishment of its life. While the sciences choke on ever
increasing flows of new detail from examining the parts of the world, not many scientists are putting
the pieces together into meaningful wholes of knowledge. We need more syntheses by more
generalists, more generalization even at the calculated risk of less accuracy.
The need is not so much for fewer specialists as it is for more generalists, not so much for less
publishing of science as for more interpretation of science where there is recognition of the value of
generalization that may be accurate enough to be of great value, while also being uncertain and
perhaps therefore inaccurate to some degree.
Conquering the tsetse fly “problem” in Africa is receiving much research. The tunnel vision
approach would focus simply on eradicating flies as a relatively simple biological problem. The
generalist might be aghast at such efforts in view of the vast ignorance of what the power to destroy
the tsetse would really accomplish through chain reaction and time. Tunnel vision sees only flies,
with perhaps another vision of people happily eating more cattle. A broader view might see over a
quarter of a continent an end result of human poverty and misery in rapidly expanding deserts that
mark the end of rich faunas and floras once containing reasonably successful people. The innocence |
of dropping sucha powerful force as easy tsetse eradication into the lap of a humanity largely blind to
the terrible powers involved has many parallels in history, where seemingly small additions to the
capability of man has resulted in world changes too big for most people to see, and too drastic for
those touched by it to understand.
Science has rarely been in the arena of public debate, expounding its discovered truths. Without
science participating in human affairs with a vigor equal to its unique capability for discovery, what
hope is there for wise choices? What hope for future life in high quality environments?
Not always do the issues have the potential inflation factor of tsetse flies or nuclear fission. But the
effects of lesser tinkerings add up, with overharvesting here, mercury waste there, disease eradication
somewhere else, each with their chain reactions. Not long ago I read an enthusiastic article froma
Canadian botany department assessing the harvest potential from the kelp growing along the British
Columbia coast. A new industry was about to be born, and the facts were proclaimed as if from the
local Chamber of Commerce. On impulse, I penned a note to the author asking to what extent the
intended harvest would be also habitat removal to the rich and specialized fauna of the kelp “forests,”
and what animal species were involved. The reply was a refreshingly candid admission that the
thought had not occurred to him. Tunnel vision was perhaps quite properly his view of the kelp beds,
8 THE CANADIAN FIELD-NATURALIST Vol. 93
but there seemed to be no other view involved. No one had considered effects and chain reactions.
I work in one of Canada’s larger museums with specialists in history, zoology, botany, ethnology,
archaeology, and other fields. When I first examined this small community of mixed disciplines, I
was impressed by the isolation from the others in which each group worked; but I became more
impressed later on those rare occasions when cross-communication between two of them triggered
an exciting and enlightening experience for both. Acommon language of biology bound together the
life sciences, and an ecological dimension common to biology and anthropology gave them common
ground from which to exchange ideas. But history was different; for traditional reasons it was unable
to comprehend either biological or ecological concepts, so was unprepared to consider them of use.
This was the most impressive museum experience ofall, and I have tried to understand it, not just for
reasons of improving a museum, but for its possible deeper meanings. The new and popular interest in
ecology has not touched the historians about me, at least not as historians. To converse with them I
must drop my ecological view of the world and lean heavily on my everyday experiences with people
interested simply in experiences with people. This sort of superficial observation of the human scene
has been man’s need and entertainment since man became man, perhaps before, and it seems that
traditional history as a discipline is largely a formalized version of this trait. A scientist might
conclude that attempts to explain history without using the knowledge of biology and ecology must
result in very limited understandings. A historian may counter that to record the passing scene 1s
enough; let others try to explain it.
The science-less view of history must be largely the view of the world as seen by most people. There
is little concern with cause and effect, the only part of the world worth noting being people. Their
successes and failures, if explained at all, result from their own interactions, chance, and the
supernatural. Modern history of course has its ecologically oriented historians, or so the literature
would suggest unless they are instead scientists trespassing in history. Nevertheless, science is not a
useful part of traditional history, not to the extent of its being a constant means of understanding the
past.
This ignoring of man’s supreme accomplishment in recent times — scientific knowledge — in
understanding the world is the standard human condition. We gladly accept the wealth and ease that
science has given us. We eagerly learn to apply the directions of science for achieving health and
comfort and riches from our environments. We spray the aphids on a bush, and thereby
unknowingly poison a hundred other kinds of creatures, including ourselves. We pour chemicals of
unknown capability into a sewer, for safety, and thereby take food in the form of fish from the
mouths of a thousand people. We remove a rich forest with machines from an endless list of
discoveries, and for quick profit burden future generations with square miles of sun-baked
wastelands. To learn to exterminate a fly may be, unknowingly, to learn how to create deserts over
half a continent. We are given power far greater than we realize, far more given to unexpected chain
reactions than we can guess, by a science not doing the important part of its job. The brilliance that
gives inventions and discoveries must come from a system discovering and disseminating also some
understanding of the impact of those inventions and discoveries upon the face of Earth.
The world needs generalists in science, using new approaches in science, to discover the true
meanings of its discoveries, and to report them in useful form giving the sure, the probable, and the
possible effects of their use. Only such synthesizers, at all levels of science, can scientifically
reconstruct whole animals, rebuild whole habitats and whole landscapes, putting aggregates of
knowledge back into the public reach so that whole things, or at least meaningful parts of things, are
understood as far as men understand them. This is not easily done. That is why science fails to do
it. By the standards of accuracy demanded in most sciences these reconstructions will have some
inaccuracy. If this is unavoidable, it must become acceptable insofar as its accuracy is useful, because
there is not just a general intellectual need for non-scientists to see more sums of knowledge, there is
some urgency since our science and technology seem to have us programmed in several ways to self-
destruct.
The challenge of evolving effective methods for such a prevalent new dimension to science may
1979 EDWARDS: GUEST EDITORIAL 9
not be the main problem. Present scientific circles frequently regard as somehow second class those of
its scientists that put much effort toward being understood in lay society. It is almost as if there was a
cult for obscurity. Perhaps we have a long way to go.
Science must of course be encouraged to pursue the obscure. But let it also learn to reassemble the
parts. Biology is not presently a force in world affairs because it does not communicate well, at least
not at the level of public understanding. Economists (and merchants, and lawyers, and others) make
decisions instead. Until biologists begin to replace economists because they have a superior ability to
correct errors and to predict events (which should not be too difficult) there is not much hope for
realistic decisions in human affairs.
If life is indeed Earth’s most important attribute, biology is then the discipline best able to guide
man’s decisions concerning his (and Earth’s) welfare. If this is even partly so, we need a biology not
Just active in knowing more of less and less, but also at knowing more of less plus less. Why? So that
people may understand their world. And so that humanity may survive with dignity.
The Canadian Field-Naturalist — the Status Quo
or a New Direction?
To some of us the most significant and lasting contribution that The Ottawa Field-Naturalists’
Club has made to society is the publication of The Canadian Field- Naturalist. From almost the very
beginning, since 1880, the club has published a scientific journal and from the early years (1889
onwards) its scope has included papers on the natural history of Canada. This centennial year of the
club is a time for contemplation, reflection, and, by the express wishes of Council, a reassessment of
The Canadian Field-Naturalist’s réle as a publication of the club and its relationship to the club.
Do Problems Exist?
In general, the present members of the Council of The Ottawa Field-Naturalists’ Club are reported
to be “uncomfortable” about The Canadian Field-Naturalist. Some would prefer more papers by
amateur naturalists, more items of local interest, more club content, and less emphasis on
professional scientific papers. Some, reflecting comments made by local members, question why a
portion (40% in 1978) of their membership fees should be allotted to a publication that appears to
have minimal relevance to the club and why those members who are not interested in The Canadian
Field- Naturalist should receive it at all. Fortunately, others understand that The Canadian Field-
Naturalist is providing a much needed service to Canadian natural history. Most people agree,
however, that the club is the most appropriate publisher of The Canadian Field- Naturalist, at least
until a more suitable alternative organization is evident.
There is no doubt that the relationship between The Canadian Field- Naturalist and The Ottawa
Field-Naturalists’ Club as its publisher is a very important one. Unfortunately, however, uncertainty
regarding this relationship continues to arise from time to time. Ina way the publication ofa national
scientific journal by a basically local natural history club is an anomaly, especially as The Canadian
Field- Naturalist is recognized as the “official” publication of the Ottawa club. It is the only
publication specifically mentioned in the club’s constitution and is sent to all club members.
Until recent years many of the members of Council were senior scientists (the Ottawa area with
many federal government departments has a high concentration of scientists) with strong interests in
the journal. They recognized its broad scope and scientific significance. Today most Council
members are not professional field-research scientists, hence the composition of Council is probably
more comparable to those of other local natural history societies. Many of the professionals (mainly
field-biologists) who were once active members of the club and served on its Council are now
involved in different activities. Therefore, is it unexpected that some members of the present Council
don’t really understand The Canadian Field- Naturalist? Their uneasiness about it is evident in their
current re-evaluation of its rdle and in the stated view of the Chairman of the Publications
Committee that the club should publish a journal that is in keeping with the wishes of the
membership. But exactly what is meant by “membership”? Is it considered to be the “local”
(arbitrarily defined geographically), largely non-professional members, the people who can most
easily communicate their opinions to Council? Or does it also include, as it should in the true sense,
the local and non-local professionals; the former local members who have moved away but still want
to keep in touch with the club; the distant people who became members because they wanted to
receive The Canadian Field-Naturalist (prior to 1976 this was the only way an individual could
receive the journal); and possibly others who, although they may dwell far from Ottawa, wish for
other reasons to belong to the club? And shouldn’t the opinions of our subscribers and other readers
also be considered?
The members of Council are voted into office at the Annual Business Meeting. The slate drawn up
by a Nominating Committee each year from the local members of the club has always been acclaimed
into office by the local members who attend. To my knowledge there has not been a single club
election in recent decades. Considering that the club’s Council is assumed to represent the
10
1979 SMITH: EDITORIAL Ui
membership, one can interpret the “wishes of the membership” to be equivalent to the wishes of the
Council members after they have taken into account the expressed opinions of local members. In
forming his/her opinion will each member of Council think of his/her own personal preferences or
will he/she consider the broader outlook, recognizing The Canadian Field-Naturalist’s role in the
advancement of scientific knowledge of Canadian natural history? And is recognition being given to
all those the journal now serves beyond the interested local members, namely the subscribers,
external members, authors, and the many readers who consult it in libraries?
The major reasons for my editorial then are: first, to alert those concerned about and interested in
The Canadian Field- Naturalist to be informed of the present uneasiness of the Council and the steps
it is planning to take; second, to solicit informed input from our readers to help the Council decide
whether the status quo should be maintained or The Canadian Field- Naturalist should take a new
direction; and third, to state my personal opinion regarding the journal’s present status and réle and
my thoughts regarding its future.
Functions of The Canadian Field-Naturalist
To me The Canadian Field- Naturalist fills the important need fora national field-oriented natural
history journal publishing, and hence recording for posterity, original and significant information
with relevance to Canadian plants and animals. For some years now there has been a drop-off in club
activity in geology and palaeontology and there has been a concomitant dearth of manuscripts
submitted to The Canadian Field- Naturalist on these subjects. This is unfortunate because these
areas of natural history were once very strong ones in the club and the journal. Thus The Canadian
Field- Naturalist currently plays a role in advancing knowledge in the biological sciences as authors
record, analyze, and interpret their data. In recent years this function has become more important
because several journals formerly available to field-biologists will no longer accommodate purely
observational (non-experimental) data nor descriptive passages outlining the results of field
investigations. The Canadian Field- Naturalist, by acting as a model for content and format, also
serves to stimulate amateurs and young biologists to record their observations and to start their own
investigations leading to fruitful lifetime hobbies or careers as field-biologists.
There is no doubt for many of us that the scientific quality of The Canadian Field- Naturalist is
currently high. Otherwise it would not attract submissions from a wide variety of natural scientists,
nor would respected scientific experts serve as Associate Editors or referees, nor would the National
Research Council of Canada have supported its publication by awarding substantial grants when
funds were needed. In earlier years, however, The Canadian Field- Naturalist was neither completely
rejected nor completely accepted by the scientific community because it did not use the peer review
system and was only partially scientific. Today it is accepted as a scientific journal and is widely
distributed, reaching many lay and professional readers including those in Canadian and foreign:
libraries. It serves to varying degrees both writers and readers by permanently preserving valuable
information of national importance and by helping to build the reputation of professionals who must
publish in a refereed journal of high standards and quality.
Although from time to time The Canadian Field- Naturalist publishes significant scientific papers
concerning Ottawa biota, I agree that in general it currently does not relate very much to local
naturalists with purely local interests other than in its publication of the reports and notices of the
club. Current editorial policy does exclude papers that are of only regional or local interest, i.e., those
that are not nationally significant, but these papers are appropriate for and can be accommodated in
society newsletters and in regional or local natural history publications, many of which are of
excellent quality. If The Canadian Field- Naturalist were to publish these items, it would again be ina
gray area and eventually it would lose its current status and reputation as a primary scientific
publication.
I feel that most naturalists should find something of interest in every issue of The Canadian Field-
Naturalist although I recognize that it does not fill the needs of all the local members. But the club
does publish five times a year an excellent magazine, 7rail & Landscape, containing items of local
| THE CANADIAN FIELD-NATURALIST Vol. 93
natural history interest, matters of conservation in the Ottawa area, plus announcements of lectures,
excursions, and special meetings of the club. As far as many local members are concerned Trail &
Landscape in fact is the voice of their club, yet it has no official status as a club publication and its
continued existence is not completely assured. Certainly re-evaluation of Trail & Landscape’s status,
réle, and relationship with the club seems urgently required in the present context.
Clearly The Canadian Field-Naturalist is filling the need for a field-oriented national natural
history journal; it serves both amateur and professional naturalists and field-biologists, maintains
high scientific standing, and has assured itself of a place in history as it records the current state of the
discipline. In doing so it is also fulfilling the second objective of the club, “to encourage investigation
and publish the results of research in all fields of natural history and to diffuse information on these
fields as widely as possible.”
Are Amateur Naturalists Considered?
One of the serious concerns of the Council of The Ottawa Field-Naturalist Club is the position of
amateurs vis-a-vis The Canadian Field-Naturalist. Various members have recently expressed
particular concern that the highly professional nature of the content and the more prestigious format
of the journal have intimidated amateur naturalists and thus prevented them from submitting
manuscripts. At Council’s request a survey was conducted of two volumes from each of the last three
decades and all authors were classified as “professionals” if they had an institutional address and as
“amateurs” if they had a home address. The survey clearly indicated a gradual decade-to-decade
decrease in the number of papers by authors classed as amateurs. But these results are open to several
interpretations. The question of whether the number of submissions from amateurs with data worthy
of publication has decreased because of the journal’s content and format cannot yet be answered. I
sincerely hope, however, that no potential author has been deterred from submitting a paper that he
or she felt was of national significance.
Let us consider some possible factors that could have contributed to the results obtained in the
survey.
a) Is an institutional address an adequate criterion for identifying an author as a professional? It
must be realized that a professional in one field of science may be an amateur in another discipline
even within the biological sciences. Also many authors prefer to use an institutional address even if
the paper they submit has no relevance to their particular profession. Thus the survey could not
identify all the amateurs in the true sense of the word.
b) Are the amateurs of today sending their manuscripts to other publications that were either not in
existence or not of sufficient interest or quality for amateurs of earlier years? In general, over the last
few decades most scientific publications, including The Canadian Field- Naturalist, have changed to
become less anecdotal, more structured, and more polished. At the same time there has been a
dramatic increase in the number and quality of alternative sources of publication for natural history
notes. Some of the regional and local publications of natural history societies have changed from
mimeographed newsletters, where often virtually any material submitted was accepted, to a printed
booklet format, where raised standards are evident. The high calibre of these publications has
enhanced the image of the organizations publishing them and surely is appreciated by their members.
Trail & Landscape is an excellent example.
c) Isamateur science recognized as the initial training ground for many of those who will eventually
make their living as professional field-biologists? Perhaps these young people with their special
enthusiasm for natural history are our most inspired amateurs. Although many manuscripts
submitted to The Canadian Field- Naturalist are from authors with institutional addresses, often the
authors are really students, 1.e., advanced amateurs who are not yet professionals.
d) Are today’s amateurs motivated to make their own observations systematically and rigorously
and to write them up for publication? In the past this motivation was often supplied by interested
1979 SMITH: EDITORIAL 13
professionals who spent considerable time answering questions, identifying organisms, and
interpreting often routine natural phenomena for members of the public, including amateur
naturalists. Amongst these were occasionally people with exceptional latent talent who needed only
a bit of interest to be shown in them by such notables as Jim Baillie and A. F. Coventry in Toronto,
Clifford Carl in Victoria, Earl Godfrey in Ottawa, and many others. Although these exceptional
professionals published valuable papers on their own research, perhaps equally important although
often unsung contributions on their part were in providing encouragement to beginning naturalists
to observe and record significant data. Because of our increasingly complex society, and the drives to
establish their own reputations and empires, it is my impression that fewer and fewer professionals in
universities, museums, and government departments take time to communicate with amateurs.
Therefore, often amateurs now cannot readily discern whether their observations, sometimes just
important single observations, are of particular interest and significance. Moreover, the incentive for
amateurs to report (write up for publication) their observations may be lacking without this
professional encouragement.
é€) Dosome of the few particularly keen amateurs who want to help advance knowledge prefer to
work under the direction of a professional field-biologist rather than on their own? Certainly
Opportunities to do this are available and advertisements specifically outlining projects are fairly
widely circulated. Undoubtedly any publications resulting from this type of joint amateur-
professional collaboration bear an institutional address.
Jf) Overall, do more amateurs today pursue natural history for the particular pleasures it brings to
them rather than through a desire to carry out a particular study or to contribute to knowledge? With
the increased leisure time in our changing society as well as increased incomes, people are travelling
more extensively, not only in Canada but all over the world. They enjoy adding new species to their
life lists, and taking photographs and making sound recordings using the array of technical
equipment now available to them. The greatly increased mobility and opportunities for fleeting but
exciting glimpses of different biomes present a far different situation than in earlier years. Then
people used to be more sedentary and amateurs were more interested and persistent in devoting time
to the valuable study of a single species or a few local species. Their repeated, often tedious,
observations over long periods of time provided data that could be synthesized and interpreted. New
trends and even definite changes were uncovered. Of course, with the advancement of knowledge, it
is no longer so easy to find new fields to conquer, e.g., to describe species or habits or extensions of
ranges. Are many of today’s amateurs still willing to do potentially valuable long-term studies in the
field and to analyze and interpret their data? Or are they preoccupied with spotting rare species here
or travelling to distant lands to identify, check off, then move on to the next, rather than studying,
watching, and recording?
g) Are amateurs reluctant to send a manuscript to The Canadian Field- Naturalist because it is too
“scientific”? Those of us currently responsible for the content of the journal are anxious to have
submitted by amateur and professional naturalists and field-biologists papers on results of sound
original investigations in any field of natural history that is of significance and relevance to Canada.
The pages of the journal are open to all amateurs whose papers meet these criteria. Reluctant
amateurs should be particularly aware that important data not put on record may be lost to science.
Although we endeavor to maintain the journal as a first-class scientific one, we also emphasize that
scientific writing does not need to be, and preferably is not, pretentious but rather is simple and
straightforward. Ideally it should be clear, concise, accurate, logical, and interesting. The journal isa
broad general one and it is important that the papers we publish be understood by as many as
possible of our readers. Although we expect trained biologists to write their manuscripts in the
accepted style (and not all do), I emphasize here that we are particularly willing to help amateurs by
giving advice and guidance on writing to improve their submitted manuscripts. The major criterion
for acceptance is the importance of the information and its relevance to Canadian natural history.
14 THE CANADIAN FIELD-NATURALIST Vol. 93
In the past, amateurs played an extremely important réle in the advancement of knowledge in the
natural sciences, particularly in field biology. Are amateurs still doing so? My questions and possible
interpretations of the results of the survey regarding amateurs may help to illustrate that answers to
seemingly simple questions are often complex and involve many interrelationships.
Future Role of The Canadian Field-Naturalist
The existence of The Canadian Field-Naturalist as a primary scientific and hence archival
publication depends on several factors. These include the submission of manuscripts that, after
review by competent referees, are found acceptable for publication; the availability of sufficient
funds to cover publication and distribution costs; and the agreement of the publisher to maintain the
journal as such.
The Articles and Notes published in The Canadian Field-Naturalist are refereed and almost all
undergo minor or major revision before they are accepted. But there are also several non-refereed
pages in each issue containing general information considered to be of wide interest. Should the
number of these pages be increased? Should they contain more items of interest only to the local club
members? For instance, would items that now appear in Trail & Landscape (sent to all local members
and any other members who request it) or in The Shrike (the club birding newsletter, available only
on subscription) be suitable? If so, would these other publications still be maintained as they are
now?
At present the major expenses of the journal are met by page charges levied to authors and the
subscription fees from institutions. Individual subscribers number just over 300 but their fees plus
the portion allotted from the membership fees are not insignificant. The results of a questionnaire
circulated to the club’s membership with the renewal forms for 1974 (see editorial in Volume 89(1),
1975) indicated that most people were members in order to receive The Canadian Field- Naturalist,
but at that time only institutions, not individuals, could subscribe. Those who were club members in
1975 were given a choice starting in 1976 of maintaining their membership in the club witha portion
of their membership fees (currently 40%) going to the journal, or of becoming subscribers for exactly
the same annual fee but with the entire fee (100%) going to the journal. I wonder how many through
knowledgeable choice, apathy, reluctance to change the status quo, or perhaps lack of knowledge of
the financial implications to the journal, chose to remain members of the club. The opportunity fora
member to become a subscriber was offered only once although there is no reason anyone wishing to
do so shouldn’t change from being a member to become a subscriber at the start of any year.
Because the club does have members across Canada and elsewhere, whether in name only or not, it
considers that it speaks for a national and international membership. Although all members
potentially have votes at the Annual Business Meeting and can elect the Council and introduce new
business at the meeting, it is the local members, now mostly non-professionals, who come to the
meeting, vote, and serve on the Council. But these are the people who will control the future of The
Canadian Field- Naturalist.
The Council of The Ottawa Field-Naturalists’ Club intends to produce within a year or soa
statement regarding the desired réles for all the club’s publications and their relationship to the club.
Thus the specific positions of Trail & Landscape and The Shrike as well that of The Canadian Field-
Naturalist will be clarified and the réle of each as a club publication defined. Because this is a
complex matter, the Council executive has assured me that any changes in direction will be
implemented slowly. Therefore, after reading the background material in this editorial, but not being
constrained by it to matters covered therein if there are other points to be brought out, interested
persons are invited to state their views. I hope that favorable as well as critical comments, if justified,
will be made because it is important to reaffirm where the journal is performing satisfactorily as well
as where it may need modifying. Subscribers have one way to influence the journal directly, and that
is by maintaining or discontinuing their subscriptions, but they can also express their views regarding
The Canadian Field Naturalist’s role as a publication of The Ottawa Field Naturalists’ Club. I hope
1979 SMITH: EDITORIAL 15
all interested readers will put forth their thoughts because there is still time to have input into the
deliberations of Council on this matter. Indeed, I invite them to do so by writing directly to the club
(Box 3264, Station C, Ottawa, Ontario K1Y 4J5); carbon copies to the Editor would be appreciated.
More formal comments as Letters to the Editor for publication are also solicited.
The 100-year history of the club and the journal is perhaps unique and it is desirable that the two
maintain a good relationship. Although the Editor and the editorial board are currently responsible
for the editorial policy and content of the journal, ultimately it is the Council’s decision that will
shape the future of The Canadian Field-Naturalist.
This editorial is a distillation of many hours of discussion and analysis of the past and the current
situation as interpreted by my husband and me. Therefore, by expressing our concerns, I hope that
the opinions solicited from readers of all categories will help to influence the outcome of the current
re-evaluation of the rdle of this journal. Furthermore, I hope the Council’s written statement will
solve the misunderstandings and misinterpretations, calm the Council’s current uneasiness, clarify
the club’s policy about its publications, and strengthen the position of The Canadian Field-
Naturalist by formal reaffirmation of its present rdle as a primary scientific natural history
publication published in Canada and relevant to Canada.
LORRAINE C. SMITH
Editor
Demographic and Dietary Responses of Red-tailed
Hawks during a Snowshoe Hare Fluctuation
ROBERT S. ADAMCIK,! ARLEN W. TODD,? and LLOYD B. KEITH!
'Department of Wildlife Ecology, University of Wisconsin, Madison, Wisconsin 53706
2Department of Recreation, Parks and Wildlife, Edmonton, Alberta
Adamcik, Robert S., Arlen W. Todd, and Lloyd B. Keith. 1979. Demographic and dietary responses of Red-tailed Hawks
during a Snowshoe Hare fluctuation. Canadian Field-Naturalist 93(1): 16-27.
Red-tailed Hawks (Buteo jamaicensis) responded functionally but not numerically to a cyclic fluctuation of Snowshoe Hares
(Lepus americanus) near Rochester, Alberta. During 1966-1975, 89% of the resident pairs laid eggs, a nesting density of | pair
per 8.3 km?. Mean clutch size (overall 2.2) ranged between years from 1.7 to 2.6, partly in response to prey-density changes.
Mean hatching dates varied from 30 May to 10 June. Marked differences in annual productivity, 0.28 to 1.90 fledged young
per breeding pair, primarily reflected nestling mortality. About 50% of all nestling losses were associated with food shortage.
Seventy percent of annual variation in mortality through age 4 wk was jointly attributable to the frequency of rain and the
weight of food brought to nestlings. The latter varied directly with Snowshoe Hare densities. The stationary population of
breeding adult redtails contrasted with the numerically and reproductively cyclic horned owls (Bubo virginianus), but
resembled redtail and other temperate-zone raptor populations further south. Food reductions during hare declines in the
boreal forest ecosystem do not seem to be critical to survival of redtail nestlings unless combined with above-average rainfall.
Key Words: Red-tailed Hawk, Buteo jamaicensis, Snowshoe Hare, Lepus americanus, population dynamics, cyclic
fluctuations.
A study of Red-tailed Hawks (Buteo jamai-
censis) was conducted in central Alberta during
1966-1975. Its objective was to examine changes
in raptor demography, food habits, and pred-
atory-prey interactions during one complete
population cycle of the Snowshoe Hare (Lepus
americanus). Early results of this work were
reported by Meslow and Keith (1966) and
Luttich et al. (1970, 1971). McInvaille and Keith
(1974) summarized findings through 1971, the
year of peak spring densities of hares on the
study area; they examined intraspecific relation-
ships among redtails and interspecific relation-
ships between redtails and Great Horned Owls
(Bubo virginianus). They also assessed the effect
of increasing hare densities on redtail breeding
biology, and estimated rates of predation on
prey populations. The most recent paper, that of
Keith et al. (1977), examined the role of Red-
tailed Hawks and other predators in the cyclic
fluctuations of Snowshoe Hares and Ruffed
Grouse (Bonasa umbellus).
The hare population at Rochester declined
after spring 1971, reaching the lowest point in its
cycle by spring 1975. Over those 4 yr, significant
changes occurred in reproduction, nestling sur-
vival, and food habits of redtails. This paper
describes such changes and examines their
relationship to prey densities. It also presents a
general analysis of redtail productivity since
1966.
Methods
Study Area
The 162-km? study area (130 km?in 1966) was
a block of mixed agricultural and forested land
near Rochester, Alberta, about 100 km N of
Edmonton. Luttich et al. (1971) and Rusch etal.
(1972) described the vegetation, topography,
and land-use practices on the area; McInvaille
and Keith (1974) classified the major habitats
and outlined their distribution as of 1971.
Except for secondary growth in fire-killed areas,
and some further clearing of forested land for
pasture, the study area changed little over the
next 4 yr.
The breeding season for redtails in central
Alberta is early April through July. Mean
temperature at Rochester during this period is
12°C, with average minimum and maximum
temperatures of 6°C and 18°C, respectively.
Temperatures near freezing have been recorded
in all summer months; and during extended
periods of cloudy or rainy weather in July and
1979 ADAMCIK ET AL.: RED-TAILED HAWKS AND SNOWSHOE HARES 17
TABLE !—Summary of weather data during 10 breeding seasons of the Red-tailed Hawk near Rochester, Alberta!
Periods in nesting cycle and types
of weather data 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 Mean
Courtship
Days in period? 26 31 31 Ds 23 DY 36 31 22 24 28
Mean daily temperature (°C) -l 2 4 9 5 5 2 3 6 5 4
Days with precipitation (%) 31 19 29 52 9 15 3] 29 32 46 29
Mean daily precipitation (mm) het Ost Ox WA Oa 0.8 AO Oss 1.1 O09)
Mean snow depth (cm) 7 25 | 2 9 25 24 6 23 8 13
Days with snow on ground (%) 65 97 29 40 52 78 83 58 82 46 63
Incubation
Days in period 32 32 32 32 32 sy) 32 32 32 32 32
Mean daily temperature (°C) 12 12 10 11 10 13 12 1] 9 11 1]
Days with precipitation (%) 44 25 34 34 22 22 DD) 38 50 53 34
Mean daily precipitation (mm) 2.1 1.2 0.8 Ie 0.4 1.0 0.9 2.1 0.7 ES) 1.2
Nestling (pre-tethering)
Days in period 28 28 23 28 26 30 28 34 24 27 28
Mean daily temperature (°C) 13 14 14 14 17 13 14 15 15 13 14
Days with precipitation (%) 39 43 57 32 39 57 50 53 42 70 48
Mean daily precipitation (mm) 1.1 0.9 1.9 ha eal 4.2 SHO AN (0) O) eles 2.6
Tethering
Days in period 20 28 27 22 28 21 23 16 31 22 2
Mean daily temperature (°C) 17 18 16 16 16 15 13 15 15 19 16
Days with precipitation (%) 60 43 67 55 64 67 61 69 74 55 62
Mean daily precipitation (mm) a) WHO Se) 4.2 6.7 5.0 IES 16 4.8 305) 3,5)
'Weather data were collected at the Meanook Meteorological Observatory, 26 km N of Rochester.
Total days in each period were determined as follows: courtship—date of first observation of a red-tail on the study area to mean date of start of
incubation; incuoation—mean date of start of incubation to mean date of hatch (32 d); nestling—mean date of hatch to mean date of tethering:
tethering—mean date of tethering to mean date of release.
August, daytime maximums around 7°C are
not uncommon. Although occasional snow
flurries occur as late as mid-May, most preci-
pitation from late April to August falls as rain.
Total spring-summer (15 August — 31 August)
rainfall is variable, ranging from 11 to 44cm
during our study. Heavy showers and rainstorms
are common, but precipitation more often
occurs as light rain over several days.
Demographic Methods
Methods of censusing raptors, tethering nest-
lings, and estimating prey densities were iden-
tical to those described by McInvaille and Keith
(1974). Snowshoe Hare and Ruffed Grouse
population estimates were revised slightly after
additional analyses. We expanded the small-
mammal census to include a spring (April-May)
index for 1973-1975, and determined summer
(June-July) densities of juvenile Richardson’s
Ground Squirrels (Spermophilus richardsonii)
on two study areas beginning in 1969. Waterfowl
data used in the present paper were based on
counts along a 0.4-km-wide aerial transect
between the towns of Cold Lake and Swan Hills.
This transect passed about 11 km N of Roches-
ter. Weather data were collected at the Meanook
Meteorological Observatory, 26km N _ of
Rochester.
Examination of Weather Data
To analyze relationships between weather and
productivity of redtails, we arbitrarily divided
their reproductive cycle at Rochester into four
stages: courtship, incubation, nestling (pre-
tethering), and tethering (Table 1). The court-
ship stage began with the first date of observa-
tion on the study area each spring. The tethering
stage ended with release of the last young we had
tethered to obtain food data. Mean dates of first
observation, start of incubation, hatch, tethering
of nestlings, and release of fledged young during
1966-75 were 5 April, 3 May, 4 June, 2 July, and
25 July, respectively.
Within the courtship stage we examined five
weather variables: mean daily temperature,
percent days with precipitation, mean daily
precipitation, mean snow depth, and percent
18 THE CANADIAN FIELD-NATURALIST
days with snow on the ground. The last two
variables were irrelevant in the three later stages
of the reproductive cycle.
Results
Prey Populations
Spring densities of adult Snowshoe Hares
peaked in 1971 at 510 per 100 ha of habitat, and
thereafter declined to only 1% of peak numbers
by 1975 (Table 2). Because the birth rate was
higher in 1970 than in 1971, young hares were
more numerous in 1970, and the total spring-
summer (May-July) population was conse-
quently higher that year. As with adults, total
hare numbers fell markedly after 1971 to about
1% of peak numbers by 1975.
Richardson’s Ground Squirrel populations
rose between 1968 and 1970, fell to about 25% of
their maximum by 1974, and increased again in
1975. During the highest year, adult densities
averaged 988 per 100 ha of habitat (closely
grazed pastures) in May, and juveniles averaged
3350 per 100 ha in June (Table 2).
Populations of voles (Microtus pennsylvan-
icus) continued to fluctuate during 1971-1975,
as in previous years. amplitudes of change
between high and low populations were 7- to 32-
fold in late summer trap-night indices. Lowest
numbers occurred in August 1968, 1971, and
1975. May populations were lower in 1973 and
1975 than in 1974 (Table 2).
Ruffed Grouse were most abundant at 61 per
100 ha of upland forest in May 1968. A major
decline took place between the springs of 1970
and 1972, as numbers fell 65% to 16 per 100 ha.
The population remained low through 1974,
then more than doubled by May 1975.
Sharp-tailed Grouse (Pedioecetes phasian-
ellus) increased from 1966 to 1970, but
decreased abruptly within the next year and were
scarce thereafter (Table 2). Waterfowl trend data
for 1971-1975 indicated a general decline of
about 50%.
Red-tailed Hawk Numbers, Reproduction,
and Nestling Mortality
During 1972-1975, the Red-tailed Hawk
population on our study area maintained the
approximate numerical stability of earlier years
(Table 3). Total individuals varied from 53 in
1967 to 39 in 1975, while breeding pairs ranged
Vol. 93
from 24 (1 per 6.8 km2) in 1966 to 16 (1 per 10.1
km2) in 1973. There was no significant difference
between years in portion of the population
paired (mean 94%), the proportion breeding
(mean 84%), or the proportion of pairs breeding
(mean 89%).
Mean hatching dates differed significantly
(P <0.05) between years (Figure 1), being latest
in 1967 (10 June) and earliest and most variable
in 1972 and 1973 (2 June and 30 May,
respectively).
Significant (P < 0.05) annual variation also
occurred in mean clutch size and mean brood
size during 1966-1975. Mean clutch sizes were
largest in 1970 and 1972 (2.6) and smallest in
1966 (1.7) and 1975 (1.9). Egg counts were made
only once per nest, ranged over the incubation
period, and included nests off the study area
(Table 3, footnote 2).
There was a marked change in the causes and
rates of nestling mortality among redtails during
1972-1975 compared with earlier years (Table
3). Losses attributable largely to horned owls
averaged 43% in 1968-1969 (Luttich et al. 1971),
but dropped to 14% in 1970-1971, as sharply
rising hare densities apparently buffered such
predation (McInvaille and Keith 1974). Preda-
tion by owls recurred in 1972 coincident with a
major decline in hares. During 1973-1975, we
recorded no horned owl predation on nestling
redtails, as Snowshoe Hares became scarce and
owl numbers declined through egress and non-
breeding (Adamcik et al. 1978). But total
nestling mortality on the study area rose from
37% in 1972 to a mean of 66% during 1973-1975.
Almost all of such losses occurred before
tethering (i.e., with 3-4 wk after hatching) in
1973-1975, whereas only about half had occur-
red by that time in earlier years.
Of the 30 young lost prior to tethering during
1966-1972, 18 disappeared completely, and the
remains of 12 (40%) were found in or below the
nest. Four of the latter had been cannibalized by
siblings.
In 1973, 8 of 18 (44%) dead young were found
in or below the nest, and 6 of these had been
cannibalized. In 1974, 4 of 14 dead young
starved in the nest, 2 disappeared, | was
cannibalized, and 7 (50%) died after falling from
the nest. Three of the latter seven birds survived
their initial fall and were replaced in the nest; two
19
ADAMCIK ET AL.. RED-TAILED HAWKS AND SNOWSHOE HARES
1979
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20 THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 3—Some population and productivity statistics for a Red-tailed Hawk population on a 162-km? study area near
Rochester, Alberta
1966 1967 1968 1969 1970 1971 1972 1973 1974 1975
Population statistics
Breeding pairs! 24 20 19 18 19 19 21 16 17 18
Non-breeding pairs ? 3 3 3 2 | 2 4 2 1
Resident singles ? 7 4 0 | 2 2 4 2 1
Productivity statistics?
Mean clutch size NWS 2.00 AMT 2.14 2.56 2.28 2.61 2.00 2.42 1.9
Mean brood size 1.71 2.00 2.07 2.14 yen y) DEG D3) 1.96 2.26 1.71
Nestling mortality (%)3
0-4 weeks 0 12 24(10) 11 5 11(16) 29(18) 64(45) 45(23) 72(78)
5-8 weeks 5 0 33(27) 28 13 0(38) 12( 6) 0(46) 17(22) 25(40)
Total 5 12 49(35) 36 17 11(48) 37(23) 64(70) 54(40) 79(87)
Young fledged per
breeding pair4 1.58 1.60 0.68 1.22 1.90 37) 1.29 0.63 0.94 0.28
‘Breeding pairs laid eggs; nonbreeding pairs did not.
?Mean clutch and brood sizes include nests off the 162-km2 study area, but within 40 km. There were 4, 1, 24, 10, 15,8, 15, and 16 off-study-area
clutches in 1966-1968 and 1971-1975, respectively; and 5, 5, 18,9, 23, 10, 14, and 13 broods. Total nestling mortality (%) equals 100 times the
complement of the product of the two-interval survival rates: thus for 1969 total mortality was 100 [1 — (0.89) (0.72)] = 36.
’Mortality rates shown in parentheses are for 18, 11, 24, 11, 13, and 13 off-study-area broods in 1968 and 1971-1975, respectively.
4Includes nests off the 162-km2 study area.
fell out again, and the third was cannibalized by
a sibling which later starved. In 1975, 15 of 18
(83%) dead nestlings died during a single 32-h
period of heavy rain, high wind, and low
temperatures. In an earlier year, 1972, 8 of 12
deaths had occurred within | wk of a heavy
rainstorm. The immediate causes of death were
thus notably different between years.
We determined pre-tethering losses among
nestlings in nests situated off the study area
during 6 of 10 yr (Table 3). In no single year
did such losses on and off the study area differ
significantly, but the overall averages of 44 and
32%, respectively, were different (P< 0.05). We
suspect that our more frequent intrusions ad-
versely affected brooding and feeding of young
at study area nests. On the other hand, as
discussed in a later section, mortality during the
tethering period (age 5 to 8 wk) on the study area
averaged significantly less (P <0.05) than
untethered young elsewhere (1.e., 15 vs. 30%).
The end result of these time- and area-specific
differences in mortality was a similar mean rate
of loss over the entire nestling period of about
50% on both areas
Dietary Responses to Changing Hare Densities
Food Habits
Snowshoe Hares, Richarson’s Ground
Squirrels, and waterfowl constituted between 59
and 89% of the weight of food for young Red-
tailed Hawks at Rochester each year (Table 4).
The remainder consisted of a variety of small-to
medium-sized mammals and birds, of which
Franklin’s Ground Squirrels (Spermophilus
franklinii), voles and sometimes grouse were
important.
Snowshoe Hares were the dominant food of
redtails during 1970-1973. Their peak use in
1970-1971 reflected the hawks’ strong func-
tional response to increasing hare densities
(MclInvaille and Keith 1974). During the next
two years hare densities declined sharply, but
there was a notable lag in the redtails’ functional
response to declining hare numbers. The con-
comitantly low ground squirrel population
(Table 2) may have been partly responsible for
this lag. By 1974, however, hares were near their
cyclic low, and ground squirrel densities were the
lowest we recorded; redtails then switched to
waterfowl, which became the most important
single food item, comprising 36% of their diet
Consumption of Richardson’s Ground
Squirrels was lowest (10% biomass) during 1972,
coincident with continued heavy utilization of
hares and a sharp decline in the ground squirrel
population (Tables 2 and 4). By 1975, with hare
densities at their cyclic low and ground squirrel
1979
ADAMCIK ET AL.: RED-TAILED HAWKS AND SNOWSHOE HARES 2)
20
MAY
25 3
©090000096
JUNE
20
10 IS
FiGure 1. Annual distribution of Red-tailed Hawk hatching dates on the 162-km? study area near Rochester, Alberta. Dates
of hatch of 63 to 84% (mean 72%) of study-area nests were determined yearly. Circles indicate nests first visited during
incubation and/or within | wk after hatching. Mean date is given at left.
densities up four-fold, ground squirrels
comprised 40% of the redtails’ diet.
Biomass of Prey Brought to Tethered Young
The amount of food supplied to tethered
nestlings varied significantly (P < 0.01) between
years (Table 5). During 1966-1971, the mean
daily biomass of prey per brood rose steadily
from 345 to 768 g, while that per individual
young increased more irregularly from 212 to
444 g owing to annual differences in brood size.
Between 1971 and 1973, the amount brought
daily to broods and individual young declined by
66 and 48% to means of 260 and 231 g,
respectively, and remained low through 1975.
Eighty-nine and 86% of the foregoing annual
variation for broods and individual young,
respectively, was attributable to changes in mean
daily biomass of Snowshoe Hares in the redtail
diet.
Annual utilization of hares was a direct
function of their densities: regressions of mean
daily biomass on total hare population estimates
(Table 2) yielded coefficients of determination
2D THE CANADIAN FIELD-NATURALIST Vol. 93
TABLE 4 — Spring-summer (1 June - 31 July) food habits of nestling Red-tailed Hawks near Rochester, Alberta!
Percent frequency Percent biomass
Prey species? 66) "67" 68 69 70) Jl 72 #73 74 75 66° 67 68 69-70) S/5 SZ See eS
Snowshoe Hare 2 4 7 6 4a Bh — 2 9 tr 0 8 S17 24") 25 46 2a eaesS 4 0
Richard’s
Ground
Squirrel Ai 3 DS DR BP) 2109 6G Ul 7 V4 42) 25 34 939) 38° 28) sO Sees,
Franklin’s
Ground
Squirrel 4 5 6 Al {ivr 3 3 2 2 2 5 7 7/ 5 l 3 5 4
Volesand mice 32 41 13 34 #40 30° 36 42 44 #23 5 6 DD 5 4 y) 4 6 10 4
Other mammals 7 10 16 13 I 5) 6 7 G@ DW 6 8 8 9 | 5 4 5
Total
mammals 6 2 6/7 7 87 8 6) Wl SO. 6 G © 7 9 8) 90 GG 77 54 74
Waterfowl 8 12 Wl Gemedte eo alOh BR eae ae 27a, alle Bel 5 1 205 Wee SOmen9
Ruffed Grouse 2 | Ore a 2 | l | tir One | SA ats) I Phar), am Ot 0
Sharp-tailed
Grouse l | | | I itr OR OR OY 2a De ne? | | ltr Or O- @
Unidentified
Grouse I I 0 I 0 eet OR Om OL reed 822 SO) 2a er Omer) | O-= Oo
Other birds 3 12 IS 10 © FT IG IS Io Bl IW »d Tile at RD! gD ee eo NO
Total birds 25m Di Boer WIS) 3675 29" 405535) 850372 25- <2) NN cls WS le ae Ome
Totals4 101 100 100 100 100 100 99 100 99 99 101 100 100 100 101 101 99 101 101 101
'Total number of food items was 695, 1063, 585, 563, 879, 545, 1224, 387, 811, and 299 in 1966-1975; biomass total (in kilograms) was 167.2,
255.0, 199.9, 161.6, 298.9, 215.6, 308.0, 85.8, 107.5, and 47.6, respectively.
*Prey not specifically identified in the table were “Voles and mice”: Microtus pennsylvanicus (average > 80% of small-mammal biomass).
Clethrionomys gapperi (7%), Peromyscus maniculatus (4%), Zapus hudsonius (1%), and Sorez cinereus (tr): “Other Mammals”: Ondatra
zibethicus, Tamiasciurus hudsonicus, Thomomys talpoides, Glaucomys sabrinus, Mustelafrenata, M. nivalis, M. erminea, Mephitis mephitis;
“Waterfowl”: Podiceps grisegena, Anas platyrhynchos, A. acuta, A. strepera, A. discors, A. carolinensis, A. americana, A. clypeata, Aythya
americana, A. collaris, A. affinis, Bucephala albeola, Porzana carolina, Fulica americana; and “Other birds”: Accipter cooperi, Falco
sparverius, Perdix perdix, Charadrius vociferus, Columba livia, Colaptes auratus, Sphyrapicus varius, Picoides villosus, Perisoreus
canadensis, Pica pica, Turdus migratorius, Sturnus vulgaris, Dendroica petechia, and unidentified songbirds and domestic chickens.
3Less than 0.5.
4Insects and amphibians constituted less than 1% of the total biomass in all years.
TABLE 5—Weight of prey items brought to tethered nestling Red-tailed Hawks near Rochester, Alberta!
1966 1967 1968 1969 1970 1971 1972 1973 1974 1975
Mean daily biomass (g)
of food per brood
Snowshoe Hare 28 Wi 121 122 289 399 229 101 14 0
Other prey species 317 375 383 457 340 369 280 159 331 316
Total (+ SE) 345 +30 452445 504+76 579+114 629+66 768 +80 5S09+53 260423 345+55 316+48
Mean daily biomass (g)
of food per individual
Snowshoe Hare 17 47 92 68 174 231 170 90 10 0
Other prey species 195 229 292 257 204 213 207 141 228 173
Total (+ SE) DIDS AN6 2762 30) S84ee al 325 22 Sl, S78 245) 7444 se 38) 3772 83" 23 sa DRS eee OMe laws
1The number of tethered broods was 14, 15, 13, 11, 17, 12, 17,7, 7, and 3 during 1966-1975, respectively. Prey biomass totals during 1966-1971 have been
recalculated since the study by McInvaille and Keith (1974), using better estimates of mean prey weights; trends remained the same as reported in that
earlier paper.
1979
(r2) of 0.89 and 0.79 for broods and individuals.
The above relationships reflect both the hare’s
periodic domination of the prey base, and the
redtail’s strong functional response to fluc-
tuating hare densities.
Nest Distribution
Using Clark and Evans’ (1954) _nearest-
neighbor test (see McInvaille and Keith 1974),
we tested for randomness the spacing within
annual nesting populations during 1972-1975.
As in previous years, the distribution of redtail
pairs on the study area was significantly
(P< 0.05) regular, suggesting that territoriality
continued to function as a spacing mechanism.
General Analysis of Factors Affecting
Productivity
The most notable demographic change among
Red-tailed Hawks from 1966 to 1975 was the
lower fledging rate during the last 3 yr: viz., 0.62
young per breeding pair in 1973-1975 vs. 1.38 in
1966-1972. Although the main immediate
reason for this decline was increased nestling
mortality (Table 3), there were significant
annual variations in mean date of nest initiation,
and mean clutch and brood size, which might
also have affected productivity. We therefore
examined each component of productivity to
determine what affected it and whether it
interacted importantly with others to influence
rates of fledging.
We considered the following demographic
variables: (1) mean date of nest initiation
(= start of incubation); (2) mean clutch and
brood size; (3) hatching success; (4) nestling
mortality to age 3-4 wk (pre-tethering); and (5)
nestling mortality from tethering to age 7-8 wk.
Where appropriate, and where data were
adequate, we examined each of the above in
relation to weather; prey density and/or biomass
brought to young; and frequency of observer
visits to nest sites during courtship, incubation,
nestling, and tethering periods.
Mean Date of Nest Initiation
Mean date of nest initiation was analyzed in
relation to weather factors and prey population
levels through simple and multiple regression.
Annual variation in mean date of nest initiation
was not significantly related to percent days with
precipitation, average daily precipitation,
average snow depth, percent days with snow on
ADAMCIK ET AL.: RED-TAILED HAWKS AND SNOWSHOE HARES 23
the ground, or mean daily temperature during
the courtship period. Nor was there any
significant relationship between annual dates of
nest initiation and spring densities of Snowshoe
Hares, Richardson’s Ground Squirrels, or
previous-fall densities of mice and voles. Dates
of nest initiation were likewise unrelated to
various combinations of prey population indices
and weather data.
Clutch and Brood Size, Hatching Success,
and Nest Desertion
We initially suspected that weather conditions
during courtship affected availability of prey
and/or redtail hunting activity, and thus might
influence clutch size. Annual variations in mean
clutch and brood sizes, however, were unrelated
to weather at Rochester. Years with largest
clutches (1970, 1972, and 1974) had a broad
spectrum of weather conditions during the
courtship period (Tables | and 3). For example,
1970 and 1972 had mean daily temperatures
during courtship of 5° vs. 2°C, rain or snow on
9 vs. 31% of the days, and snow depths of 9 vs.
24cm. In 1966, when clutches were smallest,
mean temperature and percent days with
precipation resembled conditions in 1972, while
snow depth was close to that in 1970.
Additionally, we could find no correlation
between clutch size and average daily precipi-
tation or percent days with snow on the ground.
Although our data suggest that prey
abundance may influence clutch size, the
relationship was certainly not clear-cut. The
large-clutch springs of 1970 (2.6) and 1974 (2.4)
were each associated with high vole populations
(MclInvaille and Keith 1974; Table 2); hare and >
ground squirrel densities were also high in the
first case but low in the second. In 1972, when
clutches were equally large (2.6), adult hare
densities were about 70% of those in 1970, adult
ground squirrels about 30%, and voles were little
utilized and probably scarce. The two springs in
which mean clutch size was smallest, 1966 (1.7)
and 1975 (1.9), were characterized by low hare
and vole populations; we have no information
on adult ground squirrels in 1966, but they too
were low in 1975.
It is, of course, possible that conditions on
winter ranges or during migration influence the
size of redtail clutches. There was a positive and
nearly significant correlation (r = 0.60, P = 0.07)
24 THE CANADIAN FIELD-NATURALIST
between annual dates of first arrival at
Rochester and mean dates of egg laying, but
mean clutch sizes were largely unrelated to the
latter (r = -0.34, P > 0.25). The cause of major
annual variations in mean clutch size probably
resided on the breeding grounds since largest
clutches occurred coincidently during 1970 and
1972 among redtails and non-migratory Great
Horned Owls at Rochester (Adamcik et al.
1978).
In calculating rates of hatching and nest
desertion prior to hatching, we excluded nest
desertions attributable to climbing at the onset
of laying or incubation. The significant between-
year difference in hatching and desertion rates
was entirely owing to data from 1968. In May, a
major fire swept through part of the study area
and likely prompted four nest desertions. Only
50% of all eggs hatched that year, and 6 of 19
(32%) active nests were abandoned prior to
hatching. During the other nine years, hatching
success averaged 92% (range 79 to 100%) and
nest desertion only 6% (range 0 to 17%).
Pre-tethering Mortality
Pre-tethering losses of nestlings varied
between years from 0 to 75% (Table 3). We
analyzed such mortality through stepwise
regression, in which the initial independent
variables were mean daily temperature, percent
days with precipitation, mean daily precipi-
tation, mean numbers of days between our visits
to nests, and mean prey biomass brought daily to
tethered broods (expressed both as biomass per
brood and per nestling). All statistics except the
last were obtained during the pre-tethering
period. The amount of food brought to young
was measured during the tethering period, but
we considered it also indicative of relative
amounts provided before tethering. We chose a
discriminant probability level of 10%, and
constructed two separate models, one incor-
porating mean daily biomass of prey per
tethered brood, the other per individual.
The above described regression analysis
discarded mean daily temperature, mean daily
precipitation, and our frequency of visits during
the nestling stage from those models which best
accounted for the annual variance in _pre-
tethering mortality. The resulting multiple
regression equations and coefficients of
determination were these:
Vol. 93
(1) Y; = 3.88 + 1.36A; — 0.14B;
r = 0:67; FQ,7) = 7, Bs
(2) Y,; = 8.49 + 1.25A; - 0.09C,
r =0.73; F2,7) =9:49, P1005
where Y; = estimate of pre-tethering mortality
in year 1;
Ai; = percent days with precipitation
during the nestling stage in year 1;
B; = mean biomass (g) of prey brought
daily to individual tethered young
in year 1;
C, = mean biomass (g) of prey brought
daily to tethered broods in year 1.
These models leave approximately 30% of the
variance in early nestling mortality unaccounted
for. We mentioned previously that Luttich et al.
(1971) attributed higher tethering-period
mortality in 1968 and 1969 to horned owl
predation, and MclInvaille and Keith (1974)
suggested that such predation was buffered by
the rising hare population in 1970 and 1971. A
further horned owl kill, the first since 1969,
occurred in 1972, when hares had declined
sharply and owls were still abundant. Owl
numbers fell rapidly during 1973-1975 (Adam-
cik et al. 1978), and no further predation on
tethered redtails was observed. Losses to horned
owls, either before or after tethering, would
likely be independent of weather and biomass of
prey brought to young redtails, and might thus
account for some if not most of the remaining
unexplained variance in our models.
Tethering-period Mortality
Variations in tethering-period mortality were
irregular, and independent of weather factors,
amount of food supplied by adults, or prey
population densities. This contrast with pre-
tethering mortality doubtless reflects an in-
fluence of our tethering methods, as two
important causes of pre-tethering mortality,
sibling aggression and exposure, were largely
prevented by tethering. Tethered nestlings could
not reach one another; and beginning in 1971,
tethering sites were kept dry with a suspended
plastic sheet. Thus losses of study-area young
were probably minimum estimates of normal
tethering-period mortality. This conclusion is
supported by the fact that in 2 of 6 yr
tethering-period mortality was significantly
IDS)
lower (P< 0.05) on the study area than off
(Table 3); and in at least 1 of the remaining years
(1968), the main mortality factor was owl
predation, which tethering did not prevent.
Discussion
Comparative Demography
Reported densities of nesting redtails have
ranged widely from | pair per 1.3 km? in
California (Fitch et al. 1946) to | pair per
39.5 km? in Utah (Smith and Murphy 1973). The
mean of | nesting pair per 8.3 km? at Rochester
over 10 yr was near that of | pair per 11.3 km?
reported in nine studies, including the above
two, during 18 yr (Craighead and Craighead
1956; Gates 1972; Hagar 1957; Johnson 1975;
Orians and Kuhlman 1956; Seidensticker and
Reynolds 1971).
Eleven percent of 188 pairs at Rochester did
not lay eggs during 9 yr, 1967-1975. This was
close to the 14% of 278 pairs reported on six
other study areas during 13 yr (Craighead and
Craighead 1956; Hagar 1957; Johnson 1975;
Orians and Kuhlman 1956; Smith and Murphy
1973). Such lack of nesting applied to adults, 1.e.,
birds almost 2 yr and older (Luttich et al. 1971),
because in only two cases (Gates 1972; Luttich et
al. 1971) were yearlings (brown-tailed indi-
viduals) paired. Each of these yearlings had
mated with an adult.
The annual frequency of resident singles
varied from 0 to 13% of the total Red-tailed
Hawk population at Rochester. When concerted
efforts were made in earlier studies to obtain a
total census, unmated birds were likewise often
found, but in variable numbers (Craighead and
Craighead 1956; Fitch et al. 1946; Smith and
Murphy 1973). The percentage of yearlings
among singles apparently differs markedly
between areas: all singles were yearlings in
southern Michigan (Craighead and Craighead
1956), whereas none was at Rochester. Fitch et
al. (1946). described the harassment of single
birds by territorial pairs on a California study
area, and it seems likely that a major portion of
the yearling cohort may frequent localities where
adult breeding densities are low but food 1s
plentiful.
Henny and Wight (1972) calculated that
maintenance of redtail populations was depen-
dent upon a fledging rate of about 1.35 young
ADAMCIK ET AL.: RED-TAILED HAWKS AND SNOWSHOE HARES D5)
per breeding pair. This calculation was based on
the assumptions, among others, that the 22%
nonbreeders reported in studies published
through 1970 were yearlings, and that all adults
nested. We have noted above that non-nesting
occurs among adults, averaging 13% of 466
resident pairs. This is undoubtedly a con-
servative estimate since unpaired adults are also
frequently present. Yearlings are so rarely a part
of the paired resident population that they can
be largely ignored as potential breeders.
Reduction of adult nesting from 100 to 87%
increases the estimated required fledging rate
from about 1.35 to 1.56 young per breeding pair.
These calculations are particularly sensitive to
errors in age-specific estimates of annual
mortality. As Luttich et al. (1971) pointed out,
first-year mortality may have been overesti-
mated, thereby elevating the rate of fledging
apparently needed for population maintenance.
The stationary redtail population at Rochester,
for example, fledged a mean of 1.38 young per
breeding pair during the first 7 yr of our study.
Although we may not know precisely the
fledging rate that will balance the population,
the much lower rate that occurred during the last
3 yr (mean of 0.62) was doubtless inadequate. In
the absence of compensatory changes in survival
and/or ingress, the earliest that such decreased
productivity might have been detected by us
was in 1975; but our population data are really
insensitive in this respect, because an ap-
proximately 50% drop in fledging rate would
theoretically reduce by only 2 the number of
pairs present 2 yr later.
Henny and Wight’s (1972) survey of redtail
clutch sizes implied east-west and south-north
gradients of increase. Their mean of 2.9 for 20
nests in southern British Columbia, Alberta, and
Saskatchewan was among the highest noted.
Our data from Rochester do not fit this model.
Annual means recorded by us ranged from 1.7 in
1966 to 2.6 in 1970 and 1972 (Table 3); and the
overall average of 2.2 during 1966-1975 was
comparable to the lowest regional means shown
by Henny and Wight. Our egg counts, conducted
only once during incubation and at no consistent
stage, provided an estimate of mean clutch size
over the incubation period; consequently, they
may be biased low if partial losses or clutches
occurred before hatching. We suspect, however,
26 THE CANADIAN FIELD-NATURALIST
that most earlier studies of redtails were
similarly biased.
Factors Affecting Nestling Survival
to Age 3-4 Weeks
Fluctuations in nestling mortality influenced
annual productivity of redtails at Rochester
more than did all other demographic variables.
About 70% of the variance in pre-tethering
mortality was jointly attributable to frequency
of rainfall and amount of food provided by
parent birds. These two factors are not likely
independent, because redtail hunting activity,
and probably hunting success, were reduced by
rain. The amount of food brought to nestlings
was, of course, also affected by prey densities.
Direct adverse effects of rain on survival of
nestling redtails was earlier observed in
California (Fitch et al. 1946) and New York
(Hagar 1957), but not among Red-shouldered
Hawks (Buteo lineatus) in Maryland (Henny et
al. 1973) or Buzzards (Buteo buteo) in England
(Tubbs 1974). Several times at Rochester entire
broods were found dead, wet, and abandoned
after heavy rains.
Although it was not always possible to isolate
the individual roles of inclement weather and
low prey densities in nestling mortality, we could
at least identify situations where malnutrition
rather than exposure per se appeared mainly
responsible. We conclude that emaciation,
cannibalism, and most falls from nests were
largely symptomatic of food shortage. An
unknown but substantial portion of total
disappearances was probably also caused by
cannibalism and falls, and thus likewise
ascribable to food shortage. From the above, we
estimate that approximately 50% of pre-
tethering losses resulted directly or indirectly
from a shortage of food.
Reproductive Strategy of Northern Redtails
Luttich et al. (1971) noted that breeding
populations of arctic- and temperate-zone rap-
tors responded differently to changes in prey
densities. The former, largely dependent on
widely fluctuating microtines, are highly mobile
and annually concentrate in areas where lemmings
and voles have attained temporary abundance.
Temperate-zone raptors, on the other hand,
exploit a much more diverse and stable prey
base, and local breeding populations of these
Vol. 93
raptors are often remarkably stationary from
year to year (see recent summary by Newton
1976, p. 280). Newton (1976) concluded that,
where nest sites are not limiting, regional
differences in breeding densities of diurnal
raptors reflect differences in average levels of
prey abundance. This apparent adjustment to
the food resource that usually prevails is
sometimes inadequate, for as Craighead and
Craighead (1956, p. 226) observed, “There are
many examples of hawks and eagles returning to
previous nesting territories and attempting to
raise broods where there was not sufficient food
to support them.”
Redtail breeding populations are distributed
throughout most wooded regions of the United
States and Canada south of the tundra. As a
predator, this raptor is a generalist, taking a
great variety of prey. One might thus expect that
it seldom experiences food shortage. Our studies
at Rochester suggest, however, that this may not
be true for northern redtails occupying habitats
in which Snowshoe Hares are strongly cyclic.
We estimated earlier, for example, that
malnutrition caused about 50% of the nestling
losses within 3-4wk after hatching. We
concluded too that approximately 70% of the
annual variance in such mortality could be
attributed to differences in the biomass of food
supplied to nestlings and the frequency of rain.
The former was largely determined (r? = 0.89) by
the biomass of Snowshoe Hare, which was in
turn largely determined (r2=0.89) by hare
population densities.
Despite the fact that their prey base has a
major cyclic component, breeding populations
of northern redtails exhibit the same numerical
stability and reproductive persistence that
characterizes redtails and other temperate-zone
raptors further south. The lack of any notable
reproductive response to changing hare densities
suggests to us that reductions in food during
hare declines become critical to nestling survival
only when associated with above-average rain-
fall after hatching. The latter is a post facto
random event for which an accommodating
reproductive strategy could not likely evolve.
In contrast, Great Horned Owls at Rochester
have a breeding strategy that is clearly adapted
to the hare cycle (Adamcik et al. 1978) and
differs sharply from the evident constancy of
1979
reproduction among horned owl populations
further south. As McInvaille and Keith (1974)
noted, northern horned owls unlike redtails,
“... are year-round residents, subject to the
stresses of cold and potential food-shortage
during winter. When hares and grouse are scarce
there are few alternative and available prey
species. Most small-mammal activity is sub-
nivean, and there is a paucity of overwintering
songbirds.”
Under these circumstances, cyclic lows in hare
populations are certain to affect the owls’
physical and/or physiological condition and
hence their rate of breeding in late winter
(March). When Snowshoe Hares were scarcest,
as in 1974 and 1975, horned owls at Rochester
laid no eggs, whereas during the 1968-1972
period of moderate-to-high hare populations all
pairs produced and incubated a clutch. Such
variable reproduction combined with ap-
preciable ingress and egress generated a four-
fold change in numbers of resident pairs over the
10-yr cycle of the hare (Adamcik et al. 1978).
The population dynamics of horned owls in
the boreal forest ecosystem thus resemble those
of arctic raptors (high mobility, conspicuous
density, and reproductive changes), while
breeding populations of northern redtails retain
the characteristic stability of temperate-zone
raptors.
Acknowledgments
The financial support for this study was
provided by the University of Wisconsin,
College of Agricultural and Life Sciences; the
Research Council of Alberta; the Canadian
Wildlife Service; the National Science Founda-
tion (Grant GB-12631); and the Green Tree
Garden Club, Milwaukee.
Literature Cited
Adamick, R.S., A. W. Todd, and L.B. Keith. 1978. Demo-
graphic and dietary responses of Great Horned Owls
during a Snowshoe Hare fluctuation. Canadian Field-
Naturalist 92(2): 156-166.
Clark, P. J. and F.C. Evans. 1954. Distance to nearest
neighbor as a measure of spatial relationships in popu-
lations. Ecology 35(4): 445-453.
Craighead, J.J. and F.C. Craighead, Jr. 1956. Hawks,
owls and wildlife. The Stackpole Company, Harrisburg.
443 pp.
Fitch, H.S., F. Swenson, and D.F. Tillotson. 1946.
Behavior and food habits of the Red-tailed Hawk. Condor
48(5): 205-237.
ADAMCIK ET AL.: RED-TAILED HAWKS AND SNOWSHOE HARES Di
Gates, J.M. 1972. Red-tailed Hawk populations and
ecology in east-central Wisconsin. Wilson Bulletin
84(4): 421-433.
Hagar, D.C., Jr. 1957. Nesting populations of Red-tailed
Hawks and horned owls in central New York state. Wilson
Bulletin 69(3): 263-272.
Henny, C. J. and H.M. Wight. 1972. Population ecology
and environmental pollution: Red-tailed and Cooper’s
Hawks. /n Population ecology of migratory birds. United
States Department of the Interior, Bureau of Sport
Fisheries and Wildlife, Research Report Number 2. pp.
229-250.
Henny, C. J., F. C. Schmid, E. M. Martin, and L. L. Hood.
1973. Territorial behavior, pesticides, and the population
ecology of Red-shouldered Hawks in central Maryland,
1943-1971. Ecology 54(3): 545-555.
Johnson, S. J. 1975. Productivity of the Red-tailed Hawk
in southwestern Montana. Auk 92(4): 732-736.
Keith, L. B., A. W. Todd, C. J. Brand, R. S. Adamcik, and
D.H. Rusch. 1977. An analysis of predation during a
cyclic fluctuation of Snowshoe Hares. 13th International
Congress of Game Biologists. pp. 151-175.
Keith, L.B. and L. A. Windberg. 1978. A demographic
analysis of the Snowshoe Hare cycle. Wildlife Mono-
graphs Number 58. 70 pp.
Luttich, S.N., L. B. Keith, and J.D. Stephenson. 1971.
Population dynamics of the Red-tailed Hawk (Buteo
jJamaicensis) at Rochester, Alberta. Auk 88(1): 75-87.
Luttich, S. N., D. H. Rusch, E. C. Meslow, and L. B. Keith.
1970. Ecology of Red-tailed Hawk predation in Alberta.
Ecology 51(2): 190-203.
McInvaille, W. E. and L. B. Keith. 1974. Predator-prey
relations and breeding biology of the Great Horned Owl
and Red-tailed Hawk in central Alberta. Canadian Field-
Naturalist 88(1): 1-20.
Meslow, E. C. and L. B. Keith. 1966. Summer food habits
of Red-tailed Hawks near Rochester, Alberta. Canadian
Field-Naturalist 80(2): 98-100.
Newton, I. 1976. Population limitation in diurnal raptors.
Canadian Field-Naturalist 90(3): 274-300.
Orians, G. and F. Kuhlman. 1956. Red-tailed Hawk and
horned owl populations in Wisconsin. Condor 58(5):
371-385.
Rusch, D. H., E. C. Meslow, P. D. Doerr, and L. B. Keith.
1972. Response of Great Horned Owl populations to
changing prey densities. Journal of Wildlife Management
36(2): 282-296.
Seidensticker, J.C., IV and H.V. Reynolds, III. 1971.
The nesting, reproductive performance, and chlorinated
hydrocarbon residues in the Red-tailed Hawk and Great
Horned Owl in south-central Montana. Wilson Bulletin
83(4): 408-418.
Smith, D. G. and J. R. Murphy. 1973. Breeding ecology of
raptors in the eastern Great Basin of Utah. Brigham
Young University Science Bulletin, Biological Series
18(3): 1-76.
Tubbs, C. R. 1974. The buzzard. David and Charles Ltd.,
London. 199 pp.
Received 8 May 1978
Accepted 18 September 1978
The Oriskany Sandstone Outcrop and Associated
Natural Features, a Unique Occurrence in Canada
DIANNE FAHSELT,! PAUL MAYCOCK,? GORDON WINDER,? and CRAIG CAMPBELL4
'Department of Plant Sciences, University of Western Ontario, London, Ontario N6A 3K7
2Department of Botany, Erindale College, University of Toronto, Mississauga, Ontario LSL 1C6
3Department of Geology, University of Western Ontario, London, Ontario N6A 3K7
4421 King Street North, Waterloo, Ontario N2J 3Z4
Fahselt, D., P. Maycock, G. Winder, and C. Campbell. 1979. The Oriskany sandstone outcrop and associated natural
features, a unique occurrence in Canada. Canadian Field-Naturalist 93(1): 28-40.
An unusual type of oak-hickory forest not reported previously in Ontario has been found growing on sandstone in the
southern part of the province. The sandstone is the middle Lower Devonian Oriskany Formation (approximately 380 million
years old), a unit which crops out to no significant extent anywhere else in Canada and is found only toa very limited degree in
the subsurface. This highly fossiliferous outcrop has a total areal extent of less than 250 ha (1 mi’), most of it either under a
thin cover of sandy soil or exposed as actual outcrop. Although the site is very dry it is unusually rich in plant species
composition; at least 40 tree species grow in the area including an unusually large representation of oaks. There is an
unexpectedly large number of understorey species for a dry forest. Some species in the understorey are more northerly
elements that are quite uncommon in southern Ontario; others are southern species which are rare throughout their range in
Ontario. A number of prairie associates are present as well. Twenty-two of the plants are rare in Ontario. The site serves as
habitat for Elaphe obsoleta obsoleta(Black Rat Snake), one of the largest species of snake in Canada, whose southern Ontario
populations are declining.
Key Words: Oriskany, sandstone, oak-hickory forest, big-shell community, brachiopods, Black Rat Snake.
The objective of this paper is to outline some outcrops of Oriskany sandstone are indicated on
of the outstanding natural features of the older geological maps (Stauffer 1915; Caley
Oriskany outcrop in southern Ontario. The 1940) but just one, the largest of these, has been
biologically important aspects of the site, which — verified as Oriskany using modern petrographic
have been identified only recently, are in criteria (J. F. Cowan, 1977, unpublished data,
imminent danger of being lost owing to quarry- University of Western Ontario). Only the one
ing by the Flintkote Company of Canada. An authenticated exposure is shown on recent maps
“environmental protection area” has been selec- (Sanford 1969; Hewitt and Liberty 1972; Telford
ted by the company and will remain under 1975). This outcrop of flat-lying sediments,
company ownership, but it contains little of located about 3.0 km NE of the village of Nelles
biological significance. Quarrying of the area is Corners, Regional Municipality of Haldimand-
recommended by the Ontario Municipal Board Norfolk, Ontario (42°56’N, 79°57’W) is shown
and approved by the Ontario Ministry of as an irregularly shaped area with an average
Natural Resources. Zoning by-laws were modi- diameter of approximately 1.5 km. Air photos
fied in June 1978 to permit aggregate extraction and field observations, however, reveal that the
at the Oriskany site, although the Regional outcrop is actually even more limited than the
Municipality of Haldimand-Norfolk now desig- recent maps suggest. The greater part of the
nates it as an environmentally unique ecological formation does not occur as an actual outcrop,
area. but is covered either by the younger Bois Blanc
Formation or glacial till. Figure | shows an
Geology estimate of the areal extent of the outcrop at the
The Oriskany Formation occurs to a very — surface and in the subsurface. This sketch was
limited extent in southern Ontario but in no drawn on an air photo base using observations
other part of Canada. A number of small made by ground reconnaissance.
28
1979
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FIGURE 1. Sketch based on an air photo showing the Oriskany outcrop area near Nelles Corners, Ontario. Estimated limits
of Oriskany sandstone in the subsurface are shown by a broken line. Vertical hatching represents Oriskany overlain
by the Bois Blanc Formation; the unhatched portion represents Oriskany at the surface or directly under the soil. The
locations of study sites I-IV (Tables | and 2) are indicated, as well as the open prairie area (P).
30 THE CANADIAN FIELD-NATURALIST
There is one unmapped occurrence of authen-
tic Oriskany at “Shoap’s Farm” about 2.4 kmSE
of Springvale, Ontario (42°57’N, 80°97’W)
(Parks 1913), which consists of a vertical face
only a few metres in length. As it has no
horizontal exposure, it reveals only limited
geological information and has little influence
on native plants growing on the surface. Thus,
the small Springvale outlier is of relatively little
significance.
The Oriskany sandstone outcrop near Nelles
Corners is a flat-lying erosional remnant which
is now separated by a distance of 100 km from
the edge of the continuous Oriskany in the
subsurface south of Lake Erie (Figure 2). The
EL AnKoE
Vol. 93
type locality is at Oriskany Falls, southwest of
Utica, New York. The New York correlation
chart (Rickard 1975) shows the formation as
part of the Deerparkian Stage (or the Siegenian,
in European terminology) which is middle
Lower Devonian or in absolute terms, about 380
million years old. This is an exceptionally
coarse-grained sandstone consisting mostly of
quartz with some feldspar. The quartz grains are
well rounded and closely packed with a mini-
mum of cement, which near the top of the
formation may be calcite. The degree of cemen-
tation is variable and the rock tends to be friable
at the surface where the rock is subject to frost
action. The unit is very massive occurring in beds
ONTARIO
ISOLATED
ja EES ORNS ANY ee
WR
te
OUTCROPS
GEOGRAPHIC
EXTENT (Omir mie
ORISKANY
SANDSTONE
PENNSYLVAMA
ORISKANY
OUTCROP ar 2
FiGURE 2. Geographic extent of the Oriskany Sandstone Formation. The stippled area represents Oriskany in the subsur-
face south of the Great Lakes. Isolated erosional remnants are indicated where they crop out in New York State and
Ontario. Constructed from Rickard (1975), Kreidler (1964), and Lytle (1964).
1979
up to 100cm thick, which break out in large
blocks along vertical joint planes. The rock is
porous but rain falling on the surface runs
laterally and disappears down joints or passes
through to the underlying Bass Islands Forma-
tion. The permanent water table is 16-20 m
below the surface. The imminent quarrying
operation would be carried out above the water
table.
Although fossil preservation 1s rather unusual
in a coarse sandstone, parts of the Oriskany are
highly fossiliferous. There are internal as well as
external molds which show both interior and
exterior characters of the shells. Fossils at the
Nelles Corners site have been identified by
Stauffer (1915), Caley (1940), and Best (1953)
and include brachiopods, corals, trilobites,
gastropods, pelecypods, and miscellaneous
forms. Index fossils characteristics of the Oris-
FAHSELT ET AL.: ORISKANY SANDSTONE OUTCROP FEATURES 31
kany sandstone include Acrospirifer, Costispiri-
fer, Hipparionyx, Rensselaeria, and Oriskania
(all brachiopods) (Figures 3-7). Some genera
can be found in both North America and Europe
(e.g., Acrospirifer), and others are reported only
from eastern North America (e.g., Hip-
parionyx). The large brachiopods that dominate
the fossil fauna and are referred to as a “big shell
community” (Boucot and Johnson 1967) are not
found in any other Canadian sandstone. Of the
more than 75 fossil species reported, only about
one third occur elsewhere in Ontario or Quebec.
Unconsolidated rock materials over the out-
crop are mainly of glacial origin and are either
nonexistent or very thin. The site is therefore not
suitable for agriculture and 1s largely covered by
open woods known locally as the Clanbrassil
Forest. The biological significance of the area
was not realized until relatively recently.
FIGURES 3-7. Index fossils characteristic of the Oriskany
6, Rensselaeria; 7, Costispirifer.
Formation. 3, Hipparionyx; 4, Acrospirifer; 5, Oriskania,
By THE CANADIAN FIELD-NATURALIST
Vegetation
The first indications that the vegetation was in
any way unusual were the observations of F. S.
Cook (University of Western Ontario, personal
communication 1976) who noted the presence of
saxicolous mosses which are rarely seen in
southern Ontario because their required sub-
strate is essentially unavailable. Outcrops most
common in southern Ontario are associated with
the Niagara Escarpment and they are mainly
limestone and dolostone. These rocks differ
markedly from sandstone in their calcium
content, the pH of materials derived from them
by weathering, and their suitability for certain
kinds of plant growth.
Open oak-hickory forest growing on sand-
stone bedrock is very unusual for Ontario. In
order to develop a precise idea concerning the
structure and composition of the forest, the
vegetation was examined in detail and part of the
area was sampled quantitatively.
Methods
Five homogenous areas ranging in size be-
tween about 2 and 6 ha were selected. Four were
forested and one was an opening within the
forest. A list was compiled of all trees, shrubs,
and herbs in each forested area. Two of the
larger areas were sampled quantitatively using
the Point Quarter Method (Curtis 1959); 30
sampling points were scattered at random
throughout each site. Around each point four
trees and four saplings were randomly selected in
each of four quadrants, 1.e., the nearest speci-
mens to the center point in each quadrat were
selected. At every other point a sample of the
ground layer was taken by recording presence
(frequency) in metre-square quadrats. A number
of environmental features as well as additional
structural and compositional details of the
vegetation were recorded when the sampling
procedure was completed.
A frequency value was determined for each
tree species on the basis of its percentage
occurrence at points. These values were summed
for all tree species and the contribution of an
individual species to that sum was calculated asa
percentage (= relative frequency). Similarly, the
number of stems contributed by a species at
points of occurrence was calculated as a per-
centage of the contribution of all species at all
Vol. 93
points (= relative density). The total basal area
for one species was calculated as a percentage of
basal area for all species (= relative dominance).
The three values for a given species, relative
frequency, relative density, and relative domin-
ance, were combined to give its importance
value. This is a precise measure of the ecological
influence of a tree species in the forest.
In the forest openings lists were taken of all
vascular plants present, and general observa-
tions on community structure and environ-
mental features were recorded.
Parts of the outcrop along Townline Road
(Figure 1) were badly disturbed with old
foundations, old buildings, small abandoned
quarries, and road systems. Because much of the
associated vegetation was weedy, it was not
examined in the same way as other sites within
the general area.
Results and Discussion
Open oak-hickory communities (Figure 8) of
the type located near Nelles Corners on sand-
stone bedrock had not previously been found
during an extensive survey of forest vegetation
of southern Ontario (Maycock 1963). At least 28
different tree species are found in the stands
sampled on the Oriskany outcrop area (Table I).
This is considerable diversity for sucha poor dry
site.
At Site I (Figure 1) where the soil is very
shallow there are no clear-cut dominants but
Quercus velutina (Black Oak), Q. rubra (Red
Oak), Prunus serotina (Black Cherry), Quercus
alba (White Oak), and Carya ovata (Shagbark
Hickory) are the major tree species. The source
of names used in the text and iables is Fernald
(1950). All 28 tree species are found in Site 1, but
more than half of stand importance is accounted
for by oaks which together have an importance
sum of 163 (Table 1). In the sapling’ layer
Fraxinus americana (White Ash), Prunus sero-
tina, Amelanchier arborea (Serviceberry), and
Acer saccharum (Sugar Maple) are well repre-
sented. All of the oaks are represented to a
limited extent in the sapling layer (approxi-
mately 20% of sapling-density) but are less
important here than in the tree layer. Hickory
reproduction appears to be maintaining the
status of this species. A rich assortment of tall
shrubs is found including (listed in order of
is y Ne ; nie me j 9 @
FAHSELT ET AL.: ORISKANY SANDSTONE OUTCROP FEATURES 33
FIGURE 8. Open oak-hickory forest on sandstone at the Oriskany outcrop in southern Ontario. The best of these stands are
presently scheduled for quarrying by King Paving and Materials of Burlington, Ontario, a Division of the Flintkote
Company of Canada.
decreasing dominance) Cornus racemosa (Race-
mose Dogwood), Amelanchier arborea, Vibur-
num rafinesquianum (Downy Arrow-wood),
Prunus virginiana (Choke-cherry), and Vibur-
num acerifolium (Maple-leafed Viburnum).
Pyrus coronaria (Wild Apple) is one uncom-
mon species of note. Amelanchier arborea does
not often attain tree sizes but in this stand there
are a number of large specimens. There is also a
rich tall herb layer which includes Preridium
aquilinum (Brake), Solidago caesia (Blue-stem
Goldenrod), and Aster sagittifolius (Arrow-
leafed Aster), and a low herb layer with Carex
pensylvanica, Galium aparine (Cleavers), and
G. circaezans (Wild Licorice). In total 108
species of herbs and shrubs are present. Lichens
are uncommon in forests of the deciduous forest
regions but both lichens and mosses are locally
abundant in Site I.
Site II is immediately below Site I on a
gradual south-facing slope and also has shallow
soils. The dominant trees are Red Oak and
White Oak with a lesser representation of
Shagbark Hickory and Black Cherry. Black
Cherry, Shagbark Hickory, and Red Oak are
well represented in the sapling layer, while
Cornus racemosa and Prunus virginiana are
important elements in the shrub layer. In the
medium-height herb layer Solidago caesia and
Geranium maculatum (Wild Cranesbill) are
found, and in the low herb layer are Maian-
themum canadense (\wo-leafed Solomon’s-
seal), Carex pensylvanica, and Aster macro-
phyllus (Large-leafed Aster). This is also a dry
complex with considerable diversity including
24 tree species and 87 herbs and shrubs.
Site III is just south of Site II toward the
bottom of the gentle slope. In Site III the
34 THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE |—Tree species growing in selected stands at the Oriskany outcrop area. Locations of the four sites are shown in
Figure |. Presence is indicated by plus signs and importance values (defined in text) for the two larger sites are given
in parentheses
Species
]
Acer nigrum (Black Maple) +
Acer rubrum (Red Maple) + (4)
Acer saccharum (Sugar Maple) + (9)
Amelanchier arborea (True Serviceberry) + (20)
Carpinus caroliniana (Ironwood) + (3)
Carya cordiformis (Bitternut Hickory) +
Carya ovata (Shagbark Hickory)
Celtis occidentalis (Hackberry) +
Crataegus chrysocarpa (Hawthorn) +
Crataegus sp. (Hawthorn) ste
Fagus grandifolia (Beech) -
Fraxinus americana (White Ash) + (17)
Juglans nigra (Black Walnut) +
Juniperus virginiana (Red Cedar) +
Ostrya virginiana (Hop-hornbeam) +
Pinus strobus (White Pine) +
Populus grandidentata (Large-tooth Poplar) +
Populus tremuloides (Trembling Aspen) P
Prunus cerasus (Cherry) +
Prunus serotina (Black Cherry) tp
Pyrus coronaria (Wild Apple) at
Quercus alba (White Oak)
Quercus macrocarpa (Bur Oak) F
Quercus muehlenbergii (Chestnut Oak)
Quercus rubra (Red Oak)
Quercus velutina (Black Oak)
Tilia americana (Basswood)
Ulmus americana (White Elm) +
Total trees 28
Total herbs and shrubs 108
Soil
pH 4.6-7.0
Depth A, (cm) 6
Depth B (cm) 13
Moisture dry
Sites
I] Il] IV
+ (3)
+ (3) + +
+ (18) a
+ (3) +
+ +
+ (7)
+ (21) + +
fi
A
+
+ (1) + +
i
2
+ (15) +
+ (4) + +
+ (6) + +
+ (5)
+ (19) + +
4
+ (78) + +
+ (4)
+
+ (86) + +
se (112) 7
+ (8) + +
24 13 12
87 45 27
8.0 Usd) 6.4-7.8
8 10 10
10 48 18
dry—mesic dry—mesic mesic
unconsolidated surface soil is deeper (see Table
1) than in either Site I or I]. White Oak and
Shagbark Hickory are the two dominant trees
and scattered Black Cherry is also found. In the
sapling layer cherry and White Oak are well
represented. Prominent tall shrubs are Cornus
racemosa and Viburnum acerifolium, while
herbs include Aster macrophyllus, Carex pen-
sylvanica, Galium aparine, and Potentilla sim-
plex (Old-field Cinquefoil). In total there are 13
tree species in Site II] and 45 species of herbs and
shrubs.
Site IV, like Site III, is on deeper soil. The
dominants are Sugar Maple and Red Maple
with some White Oak, White Ash, and Red Oak.
This stand is heavily shaded and the saplings are
mostly maple. The low shrub layer is very sparse
and includes scattered Viburnum acerifolium.
The only herbs are low and include Carex
pensylvanica, C. pedunculata, and Solidago
caesia. There are 12 tree species represented in
Site IV and 27 herbs and shrubs. There are few
mosses or lichens.
In the forest openings, common dominants
are Cornus racemosa, Danthonia spicata (Pov-
erty Grass), Rhus typhina (Staghorn Sumac),
Prunus virginiana, Desmodium canadense
(Canadian Tick-trefoil), Hypericum perforatum
1979
(St. John’s-wort), Solidago canadensis (Can-
adian Goldenrod), and S. juncea (Stiff Golden-
rod). One of the openings (P in Figure 1) is of
particular interest because of its high component
of dry prairie elements, notably Lespedeza
intermedia (Intermediate Bush-clover), L. capi-
tata (Headed Bush-clover), Polygala verticillata
(Whorled Milkwort), P. pol/ygama(Polygamous
Milkwort), Desmodium paniculatum (Panicu-
late Tick-trefoil), D. rotundifolium (Round-
leafed Tick-trefoil), D. dillenii, Galium pilosum,
Linum vriginiana (Virginian flax), Andropogon
gerardi (Gerard’s Beardgrass), Monarda fistu-
losa (Wild Bergamot), Physalis heterophylla
(Variable-leafed Groundcherry), Solidago nem-
oralis (Woodland Goldenrod), Panicum lineari-
folium (Linear-leafed Panic Grass), and Aster
ericoides.
The Oriskany outcrop is considered to be a
drier-than-average site for southern Ontario. If
sites in the province were ranked along a
continuum according to water availability, the
Oriskany outcrop would be the driest of those
capable of supporting forest. The trees are rather
widely spaced approaching a condition inter-
mediate between forest and savanna. The open
condition of the canopy results in high light
intensities within the stand and permits repro-
duction of species such as Black Cherry, hickory
and oaks, as well as heavy shrub and herb layers.
Populus grandidentata (Large-toothed Pop-
lar) is one species found at the site that 1s usually
characterized by a very shallow rooting system.
Many of the dominants, however, are species
able to form long tap roots. For example
Quercus velutina, Q. alba, and Carya ovata all
produce vigorous primary roots early in devel-
opment enabling the seedlings or young saplings
to withstand drought conditions. Quercus rubra
is another species represented at the site that is
capable of producing deeply penetrating tap roots
(Fowells 1965). Pinus strobus (White Pine)
typically produces just the vestige of a tap root
but has several large laterals extending outwards
and down (Fowells 1965). White Pine root
systems have been shown to penetrate toa depth
of 4.5 m (Brown and Lacate 1961).
Many trees and shrubs are first- or second-
collection records for the Haldimand County
which close gaps in the distributional patterns
FAHSELT ET AL.: ORISKANY SANDSTONE OUTCROP FEATURES 35
for species both east and west. Thus the site has
phytogeographical significance.
In the Oriskany outcrop area, the favorable
sites with deeper soil are definitely less diverse in
tree species. On better sites more dominance is
exerted by fewer species. Greater moisture, more
favorable pH, and perhaps better nutrition
contribute to this pattern of tree occupancy.
Species diversity is much greater in sites with
shallow soils (e.g., I and II). At these sites there
are a number of major tree species but no clear
dominants, and there are far greater numbers of
shrub and herb species. Diversity is undoubtedly
related to the great variety of microhabitat
available for plants. Some rock surfaces are
exposed, others are covered with thin soil. The
soil depth varies and the shallower soils have a
low pH due to sandstone and absence of
carbonate rocks providing opportunities for
acid-loving plants, conditions which are seldom
available in southern Ontario.
The maple stand (Site IV) is on deeper soil and
is representative of a type of vegetation very
common in southern Ontario. The interesting
species of the sandstone outcrop type are absent.
The sites of most value botanically are very
heterogenous at the ground surface, 1.e., those
with shallow acidic soil over sandstone. The age
of older trees in the maple stand is approxi-
mately the same as the age of older trees in the
other forested sites studied.
The openings adjacent to the forest stands
have allowed the invasion of dry prairie ele-
ments, further adding to the diversity of the area.
The high diversity indicates that there may be
other unusual species or communities in ad-
jacent areas. For example, the Dry Lake area
nearby has an interesting assortment of thicket
and seasonal meadow communities. Quercus
bicolor (Swamp White Oak) is one uncommon
species that has been found in the lowland
fringes of Dry Lake.
One group of rare and notable plants occur-
ring in the outcrop area consists of more
northern species that are quite uncommon in
southern Ontario, although they may be wide-
spread northward, and the other is composed of
southern species that are rare throughout their
range in Ontario. Northern species such as
Hairgrass (Deschampsia flexuosa) and Running
36 THE CANADIAN FIELD-NATURALIST
Clubmoss (Lycopodium clavatum) are without
doubt present here owing to the sandstone
substrate and are otherwise quite rare in south-
ernmost Ontario. Because of its southern loca-
tion, the site also contains a number of un-
common species that are members of the
Carolinian flora of Ontario. This flora occupies
the region directly north of Lake Erie to a
sinuous line from Toronto through London to
Port Franks on Lake Huron. Examples of such
species in this area are Wild Crab (Pyrus
coronaria), Black Walnut (Jug/ans nigra), Hack-
berry (Celtis occidentalis), and Chestnut Oak
(Quercus muhlenbergii). In summary, the
unique vegetational status of the area is com-
plemented by the presence of a number of rare
and interesting species. Twenty-two of the
species found in the Oriskany outcrop area are
among those that have been listed as rare by
Argus and White (1977) (Table 2). It is most
unusual to find such a large number of rare
plants concentrated in so small an area.
Habitat for the Black Rat Snake
The presence of unusual forest associations
and unusual understorey plants provides a
particular habitat for wildlife. The most notable
animal known to inhabit the Oriskany site 1s the
Vol. 93
Black Rat Snake (Elaphe obsoleta obsoleta), the
largest species of snake in Canada. The Black
Rat Snake is considered rare, threatened, or
endangered in Ontario and Canada by several
authors (Campbell 1969; Cook 1970a; Anony-
mous 1970; Froom 1972; Stewart 1974; Parsons
1976; Cook 1977; Gregory 1977).
Observations of Elaphe obsoleta at the out-
crop were made by W. W. Judd (University of
Western Ontario) in 1976 and J. Webber
(Erindale College, University of Toronto) in
1977, and by at least four local residents during
the past two years. Judd observed a snake at
close range for about | min; Webber was able to
sketch anatomical details from a distance of 2 ft.
One local person observed many snakes (of
varying sizes) simultaneously in the spring of
NSE
It should be noted that the combination of
rocky terrain and extensive relatively undis-
turbed forested area such as that found at the
Oriskany site constitutes ideal habitat for the
Black Rat Snake. Knudsen (1955) stated that the
Black Rat Snake inhabits areas that have an
abundance of rocky areas and crevices, and from
these sites they venture out into nearby agri-
cultural lands. The snake is sometimes con-
sidered a forest species (Hay 1892; Morse 1904;
TABLE 2—Rare plants (Argus and White 1977) found at the Oriskany outcrop. Areas are those shown on Figure |
Species
Asplenium platyneuron
Panicum lanuginosum var. praecocius
Carex laxiflora var. gracillima
Disporum lanuginosum
Polygonatum biflorum
Juglans nigra
Quercus bicolor
Quercus muehlenbergii
Ranunculus hispidus
Arabis canadensis
Prunus americana
Desmodium rotundifolium
Lespedeza intermedia
Linum virginianum
Viola pedata var. lineariloba
Thaspium barbinode
Vaccinium pallidum (V. vacillans)
Asclepias exaltata
Conopholis americana
Galium pilosum
Swertia caroliniensis
Aster pilosus
Common name Area found
Ebony Spleenwort I
Woolly Panic Grass P
V
Fairy Bells II
Solomon’s Seal II
Black Walnut I
Swamp White Oak V
Chinquapin Oak II
Stiffly-hairy Buttercup I
Sickle-pod ]
Wild Plum 111
Round-leafed Tick-trefoil I
Intermediate Bush-clover Pp
Virginia Flax le
Bird-foot Violet IP
Meadow Parsnip Vv
Hillside Blueberry I
II
Squawroot I
I
American Columbo V
P
1979
Wright and Wright 1957; Fitch 1963); Wright
and Wright (1957) and Fitch (1963) both
mention occurrences associated with oak-hick-
ory forests. Nearly half of the location records
for radio-implanted specimens (Fitch 1963) were
in trees while others were in burrows, low
vegetation, or buildings. At the Oriskany site
approximately 300 ha (3 km2) including the
outcrop and immediately adjacent areas are
considered suitable habitat.
In Ontario, there are now two main isolated
ranges for the Black Rat Snake. The one in the
Kingston ~ Rideau Lakes area is maintaining
itself to some degree, but the range in south-
western Ontario is clearly discontinuous despite
maps (e.g., Conant 1975) indicating a continu-
ous range along Lake Erie. In southwestern
Ontario, there are only a few small isolated
locations which are considered by C. A. Camp-
bell to be sites of Black Rat Snake populations.
These with dates of sightings in parentheses are
(1) Essex County: Point Pelee, Pelee Island
(1977), (2) Kent and Middlesex County: Skunk’s
Misery (1968, 1974), (3) Haldimand-Norfolk
Regional Municipality: the western part of the
former Norfolk County (the main center of
abundance) (1940-1978) and other locations
including the Oriskany site (1976-1978), and (4)
Niagara Region: Fonthill in the Shorthills area
(1928-1976). Even at these locations the snakes
are not abundant; they are threatened by
agriculture and a multitude of other human
activities. Clearly, the number of large snakes in
southwestern Ontario is seriously declining, as 1s
the available habitat. There is no evidence that
the Oriskany population is continuous with
others in Ontario. Further details on the Black
Rat Snake in Ontario are in an unpublished
report by C. A. Campbell (1977) entitled “The
status of the Black Rat Snake Elaphe obsoleta
obsoleta in Ontario and particularly in the
Haldimand-Norfolk Region” (available from
C. A. Campbell or D. Fahselt).
Fitch (1963) calculated that the home range of
adults in Kansas is 12 ha for males and 9 ha for
females. In Maryland, the average home range
of E. obsoleta was estimated to be 18+ha
(Stickel and Cope 1947). Small populations
are always particularly vulnerable, so the entire
forest and outcrop complex must be maintained
in its present condition in order to maximize
FAHSELT ET AL.: ORISKANY SANDSTONE OUTCROP FEATURES 37
numbers. Restriction of habitat is listed as one of
the three major factors contributing to the
decline of Elaphe obsoleta (Cook 1970a, b).
General Discussion
Threats to the Area
As the area is unique and interesting in several
ways it is useful to consider the impact of likely
types of disturbances. First, there is abundant
evidence that fires have occurred in the past.
Second, it is obvious that exposed bedrock of
this nature is an attractive commercial source of
mineral aggregate.
(a) Fire
Certainly dry sites such as this are quite
susceptible to fire. Within the area blackened
stumps or charcoal in the soil are evident at
various locations. Though some scattered trees
are older than 90 yr, it is clear that neither the
oak nor maple forest represents an advanced
state of succession. Few of the trees are large,
most having basal areas of 516 mm2 (80 in2) or
less. Fire may be one of the factors responsible
for maintaining the oak-hickory forest in its
present secondary successional state; burning
would permit oaks, hickories, and Large-
toothed Poplar to persist and predominate.
Y oung oaks and hickories, and to a lesser extent
even mature trees, sprout vigorously after the
tops are killed back to the ground by burning
(Fowells 1965); and fire may help to perpetuate
Populus grandidentata as well as some of the
prairie elements. Periodic burning may be
partially responsible for maintaining the present
successional state of the stand and contributing
to the diversity of the interesting plant species.
Selective logging at the site may have produced a
similar effect.
(6) Commercial quarrying
Beneath the Oriskany formation at the Nelles
Corners site is approximately 15m of dolo-
stone, a calcilum-magnesium carbonate (the Bass
Islands Formation), which is desirable as aggre-
gate for paving and building. Quarrying would
remove the sandstone formation upon which the
oak-hickory forest depends. The forest could not
re-establish itself afterward on the freshly ex-
posed rock surfaces of the quarry floor because
soil chemistry and water availability would be
38 THE CANADIAN FIELD-NATURALIST
quite different. Also there would be cold air
drainage into the quarry bottom. If commercial
removal of dolostone does take place an excava-
tion with sheer vertical faces would result and
this would have adverse effects on any peripheral
remnants of forest. The edge effect associated
with a disturbance as drastic as quarrying would
likely be much more pronounced in a dry area
such as this than it would in a mesic site. The
establishment of a preserve for the fossiliferous
portion of the Oriskany outcrop would not have
a sufficiently large geographic area to support
the unique plant communities such as those in
Sites I and II, nor would it be suitably placed to
include them.
Another problem associated with a possible
quarry operation would be the effects of wind-
blown particulates. Dust reduces available light
and in some cases reacts with water to form toxic
solutions. Trees growing near a source of
carbonate dust were reported to have reduced
terminal growth (Manning 1971) or to be ina
generally poor condition (Brandt and Rhoades
1972). Particulate interference with stomatal
behavior seriously affects the diffusion resis-
tance of leaves and changes the rates of gaseous
exchange (Ricks and Williams 1974). This could
aggravate moisture stress on sites that are
exposed or dry (Smith 1974). The rates of
degradation of photosynthetic pigments in the
leaves of Quercus petraea were significantly
changed owing to particulate pollutants (Ricks
and Williams 1975) and senescence occurred
earlier at the polluted sites. Leaves with moder-
ate limestone dust deposits had a greater
incidence of fungal leaf spots (Manning 1971),
and plants dusted with cement-kiln dust were
more susceptible to fungus leaf spot disease
(Schonbeck 1960).
A numerical model which predicts the rate of
dispersion of atmospheric particulates and the
amounts of deposition in wooded areas was
described by Belot et al. (1976). A forest canopy
significantly increases the concentration of par-
ticulates deposited near the source. It is pre-
dicted that maximal deposition would occur
within | km of dust-producing activities.
Tree species respond differentially to dust
accumulation. Brandt and Rhoades (1973) have
shown that while lateral growth of Liriodendron
Vol. 93
tulipifera (Tulip Tree) was increased as a result
of deposition of dust from nearby limestone
quarries and processing plants, Quercus prinus,
Q. rubra, and Acer rubrum underwent a
reduction in lateral growth. Therefore, in a
mixed stand involving these species, importance
values would be altered with time. In fact Brandt
and Rhoades (1972) documented significant
differences in the seedling-shrub and sapling
strata between two comparable sites, one witha
heavy accumulation of limestone from quarries
and processing plants and the other a control
area with no dust accumulation. The dominant
species at the dusty site would therefore change
with continued dust accumulation. The papers
by Brandt and Rhoades indicate that it would be
difficult to maintain a natural balance among
tree species in a forest adjacent to a sustained
source of heavy limestone dust.
It cannot be imagined that quarrying would
have anything but a detrimental effect on Elaphe
obsoleta. First, the snake is susceptible to road-
kill (Fitch 1963). Second, it retreats with the
advent of disturbance to wooded areas (Morse
1904; Minton 1968). Quarrying would lead to an
obvious loss of suitable habitat.
Synopsis
We have emphasized the geological impor-
tance of the Oriskany Formation in Canada and
provided some indication of the unusual nature
of this forest tract and its richness in tree and
other plant species as well as its unusual
structural and compositional features. We have
also attempted to show its value as a habitat for
the Black Rat Snake. The Oriskany site repre-
sents the on/y example of oak-hickory forest on
sandstone in Ontario. It is decidedly unique in
representing a dry upland type of oak-hickory
forest not recognized in Ontario. The oak-
hickory type in Ontario is usually associated with
heavy clay soils which have peculiar drainage
and moisture features and never seems to
approach a classical dry oak-hickory type as
does the existing example at Nelles Corners.
This feature coupled with the large number of
unique and interesting plant occurrences, es-
pecially the unusually large number of oak
species, combines to produce a natural area of
great value and interest, and one which should
1979
be given complete protection. Preservation of
the entire area would be consistent with the
adopted policy statement of the Canadian
Institute of Forestry (Weetman 1972) that
123-410 ha (300-1000 acres) is a highly desir-
able size for forested natural areas. Interesting
natural features, geological and biological, oc-
cur throughout the Oriskany outcrop area, so it
is particularly crucial in this case that a large
preserve be set aside.
The Flintkote Company has selected an
“environmental protection area” of 19 ha (47
acres) to be located along Townline Road, the
southern boundary of the quarry site. This
selection has been approved by the Ontario
Municipal Board and by the Ontario Minister of
Natural Resources. The area does contain
features of geological interest but, because of its
very small size and the disturbances there, it is of
virtually no importance biologically.
Acknowledgements
Maycock, Winder and Fahselt acknowledge
the financial assistance of the National Research
Council of Canada.
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Stewart, Darryl. 1974. Canadian endangered species. Gage
Publishing, Toronto.
Stickel, W. H. and J. B. Cope. 1947. The home ranges and
wanderings of snakes. Copeia 1947(2): 127-136.
Telford, P.G. 1974. Dunnville area. Ontario Department
of Mines, Map P988.
Weetman, G. F. 1972. Canadian Institute of Forestry poli-
cy for selection, protection and management of natural
areas. Forestry Chronicle 48: 1-3.
Wright, A.H. and A.A. Wright. 1957. Handbook of
snakes of the United States and Canada. Volume I.
Comstock Publishing Associates, Ithaca, New York.
Received 19 June 1978
Accepted 19 October 1978
Breeding Areas and Overnight Roosting Locations
in the Northern Range of the Monarch Butterfly
(Danaus plexippus plexippus) with a Summary of
Associated Migratory Routes
FRED A. URQUHART and NORAH R. URQUHART
Life Sciences, Scarborough College, University of Toronto, West Hill, Ontario
MIC 1A4
Urquhart, F. A. and N. R. Urquhart. 1979. Breeding areas and overnight roosting locations in the northern range of the
Monarch Butterfly (Danaus plexippus plexippus) with a summary of associated migration routes. Canadian Field-
Naturalist 93(1): 41-47.
Key Words: Monarch Butterfly, breeding areas, overnight roosting locations, northern range, Lepidoptera, Danaidae.
Asa result of over 40 years of investigation dealing with the ecology of the Monarch Butterfly (Danaus plexippus plexippus).
with special reference to its migratory habits, it is now possible to outline the major breeding areas in its northern range in
Canada and to explain the absence of such areas in the western provinces. The characteristics of the overnight roosting
clusters located along the migratory routes are described and compared to those of the overwintering clusters. Asummary of
the migratory routes for these northern populations is presented, based on previous published records for North America.
For the past 41 yr, commencing in the summer
of 1937, we have been involved in a long-term
study of the biology of the Monarch Butterfly,
Danaus plexippus plexippus (Danaidae: Lepi-
doptera), with special reference to its migratory
habits. Although most of our field studies and
alar tagging (Urquhart 1941, 1960) have been
centered in Ontario, observations have been
made in all provinces from Vancouver, British
Columbia to Cape North, Nova Scotia. These
field surveys were for the purpose of investi-
gating the presence, or absence, of breeding
areas and, where adults were found, to alar tag
them for migratory studies.
It is the purpose of the present paper to
summarize our field data for the northern range
in Canada, correlating this information with the
migratory habit. Because the program has been
carried out over a long period of time it has been
possible to witness the change in the northern
range of the Monarch Butterfly and the increase
in its abundance where at one time larvae were
unknown or of rare occurrence.
Methods
Field trips were carried out in various locali-
ties in each Canadian province and notes made
concerning the presence or absence of milkweed
plants (Asclepias spp.), the source of food for the
41
larvae of the Monarch Butterfly, and of the
larvae and adults of the Monarch Butterfly.
Members of the Insect Migration Association
(IMA) sent observations concerning the pres-
ence of the larvae and adults of the Monarch
Butterfly, together with notes as to their abun-
dance in each area.
An alar-tagging program (Urquhart 1960,
1965; Urquhart and Urquhart 1976b) was
carried out along with field investigations and by
members of the IMA in order to follow the
movements of the Monarch Butterfly.
Publicity arising from the recapture of alar-
tagged specimens resulted in further observa-
tions being sent to us by interested individuals.
All observations and recapture data are on
permanent file at Scarborough College, Univer-
sity of Toronto.
Results and Discussion
Breeding Range
Field surveys carried out from 1937 to 1940
indicated that breeding areas of the Monarch
Butterfly in Ontario were mostly confined to the
lower Great Lakes regions extending from
Gananoque and Kingston in the east to
Goderich in the west and south to Leamington.
Large concentrations, owing to the presence of
42 THE CANADIAN FIELD-NATURALIST
dense growths of the Common Milkweed (Asc/e-
pias syriaca), were concentrated in the Sarnia-
Leamington, Brantford-Hamilton, Toronto-
Oshawa, and Belleville regions. Although larvae
were collected further north at Barrie and
Midland they did not occur in large numbers
because growths of the Common Milkweed were
not as dense as those found further south.
As a result of the increase in the number of
roads and highways and the clearing of forested
areas for power lines, the Common Milkweed
has gradually spread northward thus increasing
the amount of this host plant for the larvae. By
1975 records of large concentrations of both
larvae and adults were reported from various
locations along the north shore of Lake
Superior, particularly at Sudbury, Sault Ste.
Marie, and Thunder Bay. Inthe early summer of
1977 larvae were more abundant in the Sault Ste.
Marie area than in the T oronto-Oshawa area; we
had larvae mailed to our laboratory in Toronto
from Sault Ste. Marie in order to carry out
various research projects.
The reason for this difference in the numbers
of Monarch Butterfly larvae in the Lake
Superior regions in the early summer as com-
pared to the Toronto-Oshawa area can be
explained. We have previously shown (Urquhart
and Urquhart 1976d, 1977) that Monarch
Butterfly migrants travel in a northwesterly
direction in the autumnal migration and north-
easterly in the vernal. In this way the migrants
reach the north shore of Lake Superior, via
Michigan, before arriving in the Ontario penin-
sular regions. Since the Common Milkweed 1s
now abundant in the Lake Superior regions
larger populations of butterflies have occurred
here in early summer than further south.
Field surveys each summer from 1962 to 1977
from Toronto north to the northeast shore of
Georgian Bay disclosed increasing abundance of
the Common Milkweed covering hundreds of
hectares along roads, highways, and unculti-
vated fields. Also, as a result of the clearing of
forested areas, with particular reference to
power lines, ever increasing areas have been
made available for the further spread of the
larval food plant. One experimental plot of
0.5 ha located in a forested area, that had been
cleared during lumbering operations, revealed a
population of 536 larvae on 15 July 1977.
Vol. 93
For other Canadian provinces records
accumulated over the past 41 yr (including 4
yr of field investigations of 1940-1944 in the
western provinces and surveys in the eastern
provinces together with numerous reports from
various interested individuals and members of
the IMA) indicate conclusively that there are no
breeding areas in British Columbia and Alberta.
A few scattered larval populations have been
reported for southern Saskatchewan (Duval)
and southern Manitoba (Transcona, Furness).
Although there have been numerous records of
adults seen in Nova Scotia, New Brunswick,
Prince Edward Island, and occasionally in
Newfoundland, we have no records of any
breeding populations there. Three reports have
been received from Quebec (Montreal, Drum-
mondville, Quebec).
The marked differences among the population
numbers in the various provinces are due in part
to the migration routes and in part to the
distribution of the species of milkweed. Since the
migration tends to a northeast-southwest direc-
tion from the overwintering Mexican Site
(Urquhart and Urquhart 1976d) the migrants
miss the western provinces with only a few
stragglers being reported. Similarly, the eastern
provinces are outside the regular migratory
route. Although species of milkweed of the
genus Asclepias are found in all provinces,
except Prince Edward Island, Labrador, and
Newfoundland, the majority of species are found
in Ontario where Asc/epias syriaca occurs in the
greatest abundance. One species is reported for
British Columbia (A. speciosa); there are no
records from Alberta: six occur in Manitoba (A.
incarnata, A. verticillata, A. ovalifolia, A.
syriaca, A. speciosa, A. viridiflora); two occur in
Saskatchewan (A. ovalifolia, A. speciosa): 10
occur in Ontario (A. incarnata, A. verticillata,
A. tuberosa, A. exaltata, A. quadrifolia, A.
syriaca, A. sullvantii, A. purpurescens, A.
hirtella, A. veridiflora); two species occur in
Quebec (A. incarnata, A. syriaca); one species 1s
recorded from New Brunswick (A. syriaca); and
one species from Nova Scotia (A. incarnata).
From the above records of the distribution of
species of the larval food plant and the direction
of migration, it is obvious why there 1s by far the
largest population in the northern range located
in the province of Ontario.
SU)
Overnight Roosting Clusters
During the autumnal migration the Monarch
Butterflies cluster on trees of various species
along the migratory routes to remain during the
night period (Urquhart 1960). Of the species of
deciduous trees chosen Red Maple (Acer rub-
rum), Sugar Maple (Acer saccharum), Mani-
toba Maple (Acer negundo), and willow (Salix
sp.) were most common. Pine (Pinus sp.) and
spruce (Picea sp.) were most frequently chosen
since it was possible for the butterflies to obtain a
secure hold with the sickle-shaped tarsal claws
(Urquhart 1960).
During periods of calm weather individuals of
a cluster are widely spaced (Figure 1). During
stormy conditions with strong winds the
migrants cluster closer together and there are
fewer clusters on the trees. The massing together
URQUHART AND URQUHART: MONARCH BUTTERFLY 43
under conditions of strong winds has the distinct
advantage of causing the weighted branch of the
tree to sway in the wind rather than being
whipped about, which would dislodge the
migrants. The leeward side is always chosen as a
protection against the wind.
Since migrants tend to cluster on certain trees
year after year, it has been thought that perhaps
an odor of some kind may have been left the
previous year thus acting as a guide for future
generations of migrants. Numerous observa-
tions in many localities, however, have not
produced evidence to substantiate such a sug-
gestion. For example, when a particular group
of roosting trees was removed, the migrants
selected another group that had not been used
previously. As has been demonstrated, the
direction of migration, the presence of suitable
FIGURE |. Overnight roosting Monarch Butterflies during the autumnal migration showing the widely scattered nature of the
clusters.
44 THE CANADIAN FIELD-NATURALIST
trees en route located near nectar-producing
flowers such as the Canada Goldenrod (Soli-
dago canadensis) and the New England Aster
(Aster novae-anglia), and topography are the
factors determining the choice of roosting trees
(Urquhart 1960).
Although overnight clusters may occur in
numerous localities throughout southern On-
tario, they are most abundant and composed of
larger clusters along the north shores of Lake
Ontario and Lake Erie. This is owing to the
migrants’ antipathy to fly over large bodies of
water beyond sight of land; hence they move in
ever increasing numbers along the lake shores
(Urquhart 1960, 1966, 1976a).
The manner in which the migrants cluster ona
particular part of the roosting tree or trees is as
follows. The first arrival flies around the tree,
testing wind direction and a suitable site upon
which to land. Many tests are made before a final
location is chosen. Having finally landed the
butterfly opens the wings to display the more
brilliantly colored dorsal surfaces. This acts as a
beacon to others that in turn land and open their
wings. If one migrant lands too close to one
already at rest the latter snaps its wings in a
warning gesture which acts as a further stimulus
for others to land upon the particular branch.
Eventually a cluster is formed.
When the migrants reach their final over-
wintering destination in the Neovolcanic Moun-
tain site of Mexico (Urquhart 1976b; Urquhart
and Urquhart 1977), they form dense clusters
that are quite unlike those of the overnight
roosts (Figure 2). Migrants on the overnight
roosts, as mentioned previously, occur as small
scattered clusters, the individuals widely spaced;
they do not cluster on the trunks of the trees or
upon adjacent bushes or upon the ground. Inthe
overwintering clusters, the roosting trees are so
densely covered that it is not possible to see the
foliage of the trees; the trunks are also densely
covered as well as the neighboring bushes. In
some loci! the ground is also covered (Urquhart
'Site refers to the geographic location of an overwintering
site, such as the Mexican Site, the California Site, etc.; Area
refers toa particular geographic location where a number of
clusters are to be found, such as a volcanic mountain in
Mexico or the Monterey Peninsula in California; Locus (1)
refers to a particular location within an area where a cluster
exists which may change from year to year, suchas lociinan
area in Mexico or in the Monterey Peninsula.
Vol. 93
and Urquhart 1977: Urquhart and Urquhart
1976a, b, d, 1977). This difference between the
two clusterings is due to the massed millions of
migrants in the overwintering roosts as com-
pared to the smaller clusters scattered over a
wide area along the migratory routes.
Migratory Routes
The Monarch Butterflies migrate from the
breeding areas in Canada mainly to Mexico where
they overwinter in various loci in various moun-
tain areas of the Mexican Site in the Neovolcanic
Mountains, sometimes referred to as the “Cross
Range” (Urquhart and Urquhart 1979).
Migrants from west of Lake Superior move
SSW (190°): those from the north shore of Lake
Huron move SSW (200°) and those from
extreme southwestern Ontario move SSW
(215°) (Figure 3, route D). Migrants from the
rest of Ontario and western Quebec indicate two
flight patterns. The greater number move SSW
(200°) to the coast of the Gulf of Mexico and
thence westward (270°) (Figure 3, route C)
following the Gulf coast eventually towards the
overwintering site (Figure 3, route E) on a tra-
jectory SSW (195°). Others, perhaps as a result
of strong westerly winds during the flight period,
move SSE (110°) (Figure 3, route A) to the
Atlantic coast. Following the coast line SSW
(195°-210°) the majority reach the coast of the
Gulf of Mexico where, following the coast line,
they travel westward (Figure 3, route C). Others,
representing a small proportion of the migrating
population, termed an “aberrant” population
(Urquhart 1976a, b: Urquhart and Urquhart,
1976c, 1977), move down the Florida peninsula
SSE (140°), thence to Cuba and Yucatan SSW
(240°) (Figure 3, route B) or, continuing SSE
(120°-140°) become scattered over the islands of
the Antilles. The final destination of this
aberrant population is as yet unknown.
Vernal Migration
Vernal migrants leave the Mexican Site in late
February and March (Urquhart and Urquhart
1979). Mating takes place at the site, when the
clusters break up, and along the migratory
routes as far as central Texas — males rarely
proceed further. They enter the various Cana-
dian provinces, especially Ontario, commencing
in the last week of May through June and early
July. During late June and early July second-
1979 URQUHART AND URQUHART: MONARCH BUTTERFLY 45
FIGURE 2. Overwintering roosting Monarch Butterflies on a tree in the Neovolcanic Plateau Site of Mexico showing the
compact nature of the clusters completely covering the branches and trunks of one of over a thousand trees.
46 THE CANADIAN FIELD-NATURALIST Vol. 93
FIGURE 3. Migration routes of the Monarch Butterfly from the breeding areas in Canada to the overwintering site in Mexico
and the aberrant migration routes to Yucatan and the Antilles. @ breeding areas; * major overnight roosting areas;
Aoverwintering site;—sdirection of flight. A, flight pattern to the Atlantic coast: B. flight pattern through Florida to Yucatan:
C, flight pattern along the north coast of the Gulf of Mexico; D, flight pattern from the western provinces, north of Lake
Huron and southwestern Ontario; E, overwintering Mexican Site; F, possible flight direction towards the Antilles.
1979
generation adults, resulting from eggs deposited
on species of milkweed by migrating females,
enter Canada. These are readily identified by
their brighter coloration, those from the Mexi-
can Site being decidedly faded with tattered wing
margins (Urquhart 1966). First-generation
adults from breeding areas in Canada appear
from mid-July through August. A second
generation occurs in the more southern sections
from mid-August through September.
Autumnal migration commences in mid-
August, the numbers increasing through late
August and September and, under conditions of
high temperature, through October. A few
stragglers have been collected in November and
three specimens during the first week of Decem-
ber. It is highly unlikely that these late migrants
reach the overwintering sites.
Acknowledgments
Financial support for the studies of the
biology of the Monarch Butterfly has been
received from the National Research Council of
Canada; the National Geographic Society,
Committee for Research and Exploration; and
the Insect Migration Association.
Literature Cited
Urquhart, F. A. 1941. A proposed method for marking
migrant butterflies. Canadian Entomologist 1941
(February): 21-22.
Urquhart, F. A. 1960. The Monarch Butterfly. University
of Toronto Press, Toronto. 361 pp.
Urquhart, F.A. 1965. Monarch Butterfly (Danaus
plexippus) migration studies: autumnal movement.
URQUHART AND URQUHART: MONARCH BUTTERFLY 47
Proceedings of the Entomological Society of Ontario 5:
23-33.
Urquhart, F. A. 1966. A study of the migration of the Gulf
Coast populations of the Monarch Butterfly (Danaus
plexippus L.) in North America. Annales Zoologici
Fennici 3: 82-87.
Urquhart, F. A. 1976a. Migration of butterflies along the
gulf coast of northern Florida. Journal of the Lepidop-
terists’ Society 30: 59-61.
Urquhart, F. A. 1976b. Found at last: the monarch’s winter
home. National Geographic Magazine 150: 161-173.
Urquhart, F. A. and N.R. Urquhart. 1976a. Monarch
Butterfly (Danaus p. plexippus L.) overwintering popula-
tion in Mexico (Lepidoptera: Danaidae). Atalanta 7:
56-60.
Urquhart, F. A. and N.R. Urquhart. 1976b. Ecological
studies of the Monarch Butterfly (Danaus p. plexippus
L.). National Geographic Society Research Report 1976:
437-443.
Urquhart, F. A. and N.R. Urquhart. 1976c. A study of
peninsular Florida populations of the Monarch Butterfly
(Danaus p. plexippus L.; Danaidae). Journal of the
Lepidopterists’ Society 30: 73-87.
Urquhart, F. A. and N.R. Urquhart. 1976d. The over-
wintering site of the eastern population of the Monarch
Butterly (Danaus p. plexippus L., Danaidae) in southern
Mexico. Journal of the Lepidopterists’ Society 30:
153-158.
Urquhart, F. A. and N. R. Urquhart. 1977. Overwintering
areas and migratory routes of the Monarch Butterfly
(Danaus p. plexippus; Lepidoptera: Danaidae) in North
America, with special reference to the western popula-
tion. Canadian Entomologist 109: 1583-1589.
Urquhart, F. A. and N. R. Urquhart. 1979. Vernal migra-
tion of the Monarch Butterfly (Danaus p. plexippus L.;
Lepidoptera: Danaidae) in North America from the
overwintering site in the neovolcanic plateau of Mexico.
Canadian Entomologist (In press).
Received 25 May 1978
Accepted 14 August 1978
Reproductive Biology of the Big Brown Bat
(Eptesicus fuscus) in Alberta
DAVID B. SCHOWALTER and JOHN R. GUNSON
Alberta Fish and Wildlife Division, 6909-116 St., Edmonton, Alberta T6H 4P2
Schowalter, D. B. and J. R. Gunson. 1979. Reproductive biology of the Big Brown Bat (Eptesicus fuscus) in Alberta.
Canadian Field-Naturalist 93(1): 48-54.
Data on Big Brown Bat (Eptesicus fuscus) populations in Alberta were collected from 1972 to 1977 during surveys to
determine the prevalence of rabies. The 60 maternity colonies located included 56 in older buildings; many of these sites were
used as hibernacula. Most young were born during the latter part of June; parturition was estimated to extend from at least 5
June to 12 July. Fifteen of 115 pregnancies were twins. Ages of a sample of bats were determined by counts of dental annuli;
those ages generally correlated with tooth wear. Yearling females had a lower pregnancy rate than older females.
Key Words: Big Brown Bat, life history, dental aging, Alberta.
The Big Brown Bat (Eptesicus fuscus) 1s
relatively abundant and widely distributed over
North America (Hall and Kelson 1959; Barbour
and Davis 1969). Most studies of the species
have been in eastern North America (Christian
1956; Brenner 1968; Davis et al. 1968; Barbour
and Davis 1969; Kunz 1974; Mills et al. 1975)
where the usual litter size is two. Comparatively
few data are available for the species in western
North America, particularly western Canada,
where the usual litter size is evidently one
(Cockrum 1955; Christian 1956; Kunz 1974).
A bat-rabies monitoring program in Alberta
was prompted by the first diagnosis of a rabid
bat in the province in 1971, and an outbreak of
rabies among other animals in 1970 and 1971.
Epidemiological results of this program have
recently been presented by Dorward et al.
(1977); biological data on Ef. pallidus in
Alberta collected during the course of the study
are presented here.
Methods
Colonies were located through the following
means: complaints of bats relayed by govern-
mental personnel and systematic surveys during
1971 to 1977; newspaper advertisements in 1971
and 1972; results of a questionnaire mailed to
rural homeowners in 1973; and reviews of
histories of rabies-suspect bats. Collections of 10
to 20 bats for rabies testing were made at a
number of maternity roosts; larger collections
48
were made at two maternity roosts and a fall
roost, all of which were known to have harbored
rabid individuals. Bats were collected within
colonies by hand, forceps, and a modified
Constantine trap (Constantine 1958); at colony
exits by a variety of enclosure traps; and by
mist-nesting away from colonies. Colonies were
generally visited only once, although two or
three visits per year were made to one colony to
band bats. Available for analysis were 256 Big
Brown Bats submitted as rabies-suspect indi-
viduals from 1974 to 1977.
Age (adult or juvenile) as determined by
closure of the epiphyses of the fingers, and sex
were noted for each bat. Reproductive status of
adult females was recorded, as was the number
and sex of fetuses of parous individuals. Ages of
adult bats taken at two maternity roosts, a fall
roost, and a sample of rabies-suspect individuals
were determined from counts of dental annuli
(Schowalter et al. 1978). The first of January was
arbitrarily used as the date on which bats became
a year older: thus a bat classified as a l-yr-old
was taken the summer after the summer of its
birth. Tests for significance were by the simple
chi-square test.
Results and Discussion
Big Brown Bats appeared to be more abun-
dant in southern than central Alberta (Figure 1).
We discovered 32 colonies in southern Alberta
and 28 colonies in the larger area of central
1979
HINTON
Colony Size
@® 1-25
@ 26-250
ee Over 250
(0) 50 100 Km
+4
FIGURE |. Distribution of known
Alberta. Although estimates of the number of
bats were approximate, colonies were evidently
larger in the south, where the average number of
adults was 80. In central Alberta average colony
size was 44. Mist-net captures indicated that Big
Brown Bats were relatively abundant in natural
habitats along river valleys in the southern area
of the province. Much of this apparent greater
abundance in the south is related to the
occurrence of maternity roosts in the cities of
Medicine Hat and Lethbridge. We have seldom
located maternity roosts of Big Brown Bats, or
other species, in more northerly cities in Alberta.
A search of 28 old buildings in Edmonton,
mostly schools, which were similar to buildings
SCHOWALTER AND GUNSON: BIG BROWN BAT, ALBERTA 49
-—
_—_— —
= aw = =,
=-
EOMONTON
a
e e
C )
S O CAMROSE
e @ °
@
@ e®
®
e @
g
RK) CALGARY
Big Brown Bat colonies in Alberta.
frequently colonized in Medicine Hat and
Lethbridge, produced no evidence of active
maternity colonies.
A strong preference was evident for the
formation of maternity colonies in older build-
ings; 56 of 60 buildings with Big Brown Bat
maternity colonies were built prior to 1925.
Banding studies (Beer 1955; Davis et al. 1968;
Barbour and Davis 1969) have demonstrated
strong site attachment of Big Brown Bats.
Failure to colonize newer buildings may in-
dicate a decreased or stable population that is
strongly attached to roosts currently in use.
Alternatively the variable environment in older
buildings may meet the physiological needs of
50 THE CANADIAN FIELD-NATURALIST
the bats better, by permitting both summer and
winter roosting. Many nurseries were evidently
utilized as hibernacula by at least a few (and
often many) bats in Alberta, as noted by others
(Mills et al. 1975; Hitchcock 1949). Environ-
mental conditions in newer buildings are likely
to be different as a result of recent changes in
heating systems and insulation standards.
Big Brown Bat roosts were similar to those
described elsewhere (Barbour and Davis 1969;
Mills et al. 1975); those we observed were
generally cooler than Little Brown Bat (Myotis
lucifugus) maternity roosts inspected. Big
Brown Bats were extremely tolerant of light in
some roosts.
Many owners of the buildings that were used
as maternity roosts by Big Brown Bats believed
that bats were resident during the winter,
although only 19 reported having seen bats in
this period. Other buildings, particularly in the
city of Edmonton, that were not maternity
roosts were utilized by wintering bats. Our
winter captures of Big Brown Bats, and citizen
submissions of bats for rabies testing from
Edmonton, far exceeded those of summer. On
the other hand, numerous summer submissions
were made of Little Brown Bats and a few Silver-
haired Bats (Lasionycteris noctivagans) (Dor-
ward et al. 1977; this study, unpublished data).
These observations suggest that Big Brown Bats
moved into Edmonton to hibernate.
Five buildings in Edmonton and one non-
maternity roost building in Camrose and Hinton
regularly were occupied by bats in winter. As
many as six bats captured in one winter at two of
these sites were known to us; however, it is
unlikely that most bats encountered in these
buildings were reported. Winter occurrences
were frequently associated with the onset of cold
weather. In addition Big Brown Bats have been
discovered during the course of building demoli-
tion and remodelling; and in one case, loud rock
bands in a high school gymnasium were con-
sidered almost certain to cause bat activity.
As noted by Barbour and Davis (1969) E.
fuscus is extremely cold-tolerant. Two indi-
viduals were captured outside a building on 3
December 1975. Temperature at time of capture
was near —15°C, and had been even lower earlier
in the day. One bat was alert and shivering; the
other was torpid, but quickly became active
Vol. 93
when warmed. The bats had roosted near a
recently filled crack and may have been attempt-
ing to enter the building.
Timing of events at maternity roosts was
difficult to define as visits were made ir-
regularly. It was evident, however, that there was
variation between colonies, and probably be-
tween years. Bats were active at one location,
which was thought to be a hibernaculum as well,
on 8 April 1976 and 13 April 1977. The nature of
bat activity at these times was unknown as the
apparent food supply was extremely limited.
One maternity site was determined to be without
bats as late as 10 May 1977.
Roosting in four types of groups (pregnant
females, females with naked young, furred
young, and lactating females) was observed in
one maternity roost. Similar separate roosting,
although usually in only two or three types of
groups, was noted in other colonies. This
grouping created uncertainty as to the timing of
parturition, as generally not all the bats in a
roost were known to have been observed. These
groupings were thought to be related to differ-
ing thermal requirements of individuals at
different stages of reproduction and growth.
Time of parturition varied considerably both
in, and between, colonies. Fetal examination
and observation of neonates indicated that most
young were born in the latter part of June. The
earliest estimated birth date was 5 June. Near-
term pregnant females were taken as late as 12
July.
Of 114 adult females taken from maternity
roosts during late May and early June, 105 were
pregnant. Of 115 bats examined in advanced
pregnancy, 15 had twins, which corroborates
Christian’s (1956) finding that one is the usual
litter size in western North America. Sixty-two
of 84 single young had implanted in the right
horn of the uterus. Kunz (1974) also noted a
tendency towards implantation in the right horn.
Fetal sex ratio was near 50:50 (Table 1).
During 1974 and 1975 significantly more adult
females (P< 0.05) than adult males were sub-
mitted for rabies testing (Table 1). During these
two years public concern about bat rabies was
high; more than 400 bats were submitted per
year. It appeared that many normal-acting bats
were submitted as rabies suspects from mater-
nity roosts. Most bats submitted in 1976 and
1979
SCHOWALTER AND GUNSON: BIG BROWN BAT, ALBERTA 5]
TABLE |—Age-specific sex ratio in samples of Epresicus fuscus from Alberta, 1972 to 1977
Sample Adults Juveniles Fetuses
No. % Male No. % Male No. % Male
Colony survey
20 May-31 July 261 3.1 100 45.0 73 S23
1 Aug-30 Sept 69 23.2 79 26.6
Rabies suspect
1974-1975 107 36.4 46 60.9
1976-1977 58 S522 45 71.1
1977, when approximately 200 bats a year were
submitted, were either grounded or found in
other atypical situations. In those two years the
numbers of adult males and females submitted
were similar (Table 1).
Juvenile males tended to leave colonies earlier
than juvenile females during late July and
August. They occurred significantly less fre-
quently (P< 0.01) than juvenile females in our
samples from buildings during August and
September (Table 1). This earlier departure
may have subjected them to higher mortality, as
they were submitted for rabies testing more
frequently (P< 0.05) than juvenile females
(Table 1).
Results of dental aging (Figure 2) indicated a
life-expectancy similar to that found by Goeh-
ring (1972) from a 20-yr banding study in a
Minnesota hibernaculum. Our samples varied in
age structure, particularly in the proportion of I-
yr-old bats (Figure 2). One-year-olds constituted
39% of the rabies-suspect sample; the sample
was comprised of bats from the greatest variety
of situations and capture dates, and may be the
most representative of the population age
structure. It is, however, made up of bats that
died from natural or human causes. One-year-
old females from maternity colonies made up
only 16% of the sample (Figure 2). That only 15
of 31 l-yr-old rabies-suspect and survey females
were parous or had suckled young demonstrated
that many lI-yr-old females are non-parous as
noted by Christian (1956) and Barbour and
Davis (1969). Non-parous 1|-yr-old females may
tend not to roost in maternity colonies and may
be more likely to be submitted as rabies-suspect
bats than older females.
Ages of bats from Connaught School (Figure
2) have been discussed by Schowalter et al.
(1978). Younger animals appeared to be under-
represented in that sample compared to the
other samples and what would be expected from
banding results (Goehring 1972). Estimates of
the total number of adults in the Connaught
colony in June ranged from 500 to 1000,
comparable to the largest colonies noted by
Mills et al. (1975). Those authors determined
that there was an unknown density-dependent
mechanism functioning to regulate populations
in large colonies. The relatively small number of
individuals in some of the younger age-groups in
the Connaught sample suggests low survival of
young or that a large proportion of juvenile
females disperse from the colony prior to their
second year. The pregnancy rate of adult females
from Connaught, 45 of 51, while less than that of
females from other colonies (P< 0.05), is not
low enough to suggest that the regulatory
mechanism operates primarily through limiting
reproduction.
Tooth wear has been considered as a means of
aging bats (Twente 1955; Christian 1956; Stege-
man 1956). Mills et al. (1975) found a highly:
significant relationship between tooth wear and
ages of 208 Big Brown Bats banded as im-
matures. They observed wide variation in wear
rates of individual bats. One of us (JRG)
recorded tooth wear of bats in the Connaught
School sample. Although analysis was limited
by the subjective nature of the estimates of tooth
wear and the possibility of errors in aging by
annuli, there was general agreement between the
two methods (Figure 3). Tooth wear would
appear to be potentially useful as a method of
determining relative population-age structures
of Big Brown Bats in Alberta.
The possibility of a population decline and the
factors governing the growth and size of colonies
S52 THE CANADIAN FIELD-NATURALIST Vol. 93
Connaught School
Maternity Sample
n=51
Stony Plain
Maternity Sample
White Elephant
Fall sample
=43
Percent of Sample
Rabies Suspect
n= 49
12 Si waa eal 7emoie > elOm lal l2ilS ue lAghlow Onl Zale
Age Group
FIGURE 2. Distribution of ages of samples of Big Brown Bats from Alberta (determined by counts of dental annuli).
Specimens from Connaught School, Medicine Hat; Stony Plain; and White Elephant Theatre, Bow Island. Numerals
at base of bars indicate number of bats.
1979
—~-~NWAU—NWEUNUAN DWDODO,H NW EN AN © DO O
Number of Bats
EF] EI = eo es
SCHOWALTER AND GUNSON: BIG BROWN BAT, ALBERTA 53
Light wear
n=19
x=2.9
Moderate wear
n= 19
x= 5.8
Heavy wear
n= 13
x= 9.)
De Si Ae Ol 72a, LOnill 2) 1S 42 lo Onl7a18" 19-20,
Dental Age Group
FIGURE 3. Comparison of amount of tooth wear and ages, as determined by counts of dental annuli, of Big Brown Bats
from Connaught School.
in central and southern Alberta need further
investigation. Dental aging appears to offer a
powerful tool in such investigations, although
this technique requires further evaluation.
Acknowledgments
The interest and support of H. Vance and G.
Whenham of the Veterinary Services Division,
Alberta Department of Agriculture are greatly
appreciated. The Agriculture Canada Animal
Diseases Research Institute (Western) at Leth-
bridge, Alberta has provided continued co-
operation in allowing access to specimens and
records of rabies-suspect animals; W. Dorward,
J. Bradley, H. Boumans, B. Prins, and D.
Meyers of that institution contributed in various
ways to this study. Field and laboratory assis-
tance were provided by L. Harder, B. Treichel,
L. Dube, P. Cole, W. Johnson, and W.
Wynnyk. N. Previsich contributed generously of
his time and energy. We thank A. Todd and L.
Harder for comments on an earlier draft.
Literature Cited
Barbour, R. W. and W. H. Davis. 1969. Bats of America.
University Press of Kentucky, Lexington. 286 pp.
Brenner, F. J. 1968. A three-year study of two breeding
colonies of the Big Brown Bat, Eptesicus fuscus. Journal
of Mammalogy 49: 775-778.
54 THE CANADIAN FIELD-NATURALIST
Beer, J. R. 1955. Survival and movements of banded Big
Brown Bats. Journal of Mammalogy 36: 242-248.
Christian, J.J. 1956. The natural history of a summer
aggregation of the Big Brown Bat, Eptesicus fuscus
fuscus. American Midland Naturalist 55: 66-95.
Cockrum, E. L. 1955. Reproduction of North American
bats. Transactions of the Kansas Academy of Sciences
58: 487-511.
Constantine, D. G. 1958. An automatic bat-collecting de-
vice. Journal of Wildlife Management 22: 17-22.
Davis, W. H., R. W. Barbour, and M.D. Hassell. 1968.
Colonial behavior of Eptesicus fuscus. Journal of Mam-
malogy 49: 44-50.
Dorward, W.J., D.B. Schowalter, and J. R. Gunson.
1977. Prelimary studies of bat rabies in Alberta. Can-
adian Veterinary Journal 18: 341-348.
Goehring, H.H. 1972. Twenty-year study of Eptesicus
fuscus in Minnesota. Journal of Mammalogy 53: 201-
207.
Hall, E.R. and K.R. Kelson. 1959. The mammals of
North America. Volume |. Ronald Press, New York.
546 pp.
Vol. 93
Hitchcock, H. B. 1949. Hibernation of bats in southeastern
Ontario and adjacent Quebec. Canadian Field-Naturalist
63: 47-59.
Kunz, T. H. 1974. Reproduction, growth and mortality of
the vespertilionid bat Eptesicus fuscus in Kansas. Journal
of Mammalogy 55: 1-13.
Mills, R. S., G. W. Barrett, and M. P. Farrell. 1975. Pop-
ulation dynamics of the Big Brown Bat ( Eptesicus fuscus)
in southwestern Ohio. Journal of Mammalogy 56: 591—
604.
Schowalter, D. B., L. D. Harder, and B. H. Treichel. 1978.
Age composition of some vespertilionid bats as deter-
mined by dental annuli. Canadian Journal of Zoology
56: 355-358.
Stegeman, L. C. 1956. Tooth development and wear in
Myotis. Journal of Mammalogy 37: 58-63.
Twente, J. W., Jr. 1955. Aspects of population study of
cave-dwelling bats. Journal of Mammalogy 36: 379-390.
Received 14 June 1978
Accepted 18 September 1978
Wild Mallard Stocking in a Large Marsh Habitat
ROBERT O. BAILEY
Macdonald College of McGill University, Ste. Anne de Bellevue, Québec
HOA !CO
Bailey, R.O. 1979. Wild Mallard stocking in a large marsh habitat. Canadian Field-Naturalist 93(1): 55-62.
During 1971 and 1972, 1204 female and 214 male wild-strain Mallard (Anas platyrhynchos) ducklings were released on the
Delta Marsh to test Mallard stocking as a method to increase breeding populations. Hand-reared yearling hens arrived after
most of the unmarked hens in spring. Homing to the release site was observed in 26-28% of yearling hens and 53% of 2-yr-olds.
Yearling marked hens initiated nests later and were less successful than unmarked hens in producing broods. Differences
between the marked yearlings and unmarked breeders were attributed to the presence of the adult hens in the unmarked
portion of the breeding population. In view of the poor reproductive success of hand-reared birds and apparently high
potential for natural immigration and production, Mallard stocking is a questionable procedure on the Delta Marsh.
Key Words: Mallard, stocking, spring arrival, homing, nest initiation, productivity, breeding success.
Early propagation efforts involved mainly the
release of semi-domestic strains of Mallard
(Anas platyrhynchos) ducklings. Lincoln (1934),
Errington and Albert (1936), Benson (1939),
Foley (1954), Hunt et al. (1958), and Schlad weiler
and Tester (1972) concluded that releases of
game farm birds were generally unsuccessful,
because of the inability of these birds to survive
and reproduce in the wild, and because they were
extremely vulnerable to hunting. Stocking pro-
grams using wild-strain birds, however, have
been successful in establishing Mallards (Foley
et al. 1961; Lee and Kruse 1973) and Wood
Ducks (Aix sponsa) (Doty and Kruse 1972) on
vacant or understocked habitat.
In 1969 the Delta Waterfowl Research Station
began to study the potential of releasing hand-
reared wild Mallard ducklings to increase local
breeding populations. Sellers (1973) made sev-
eral releases into pothole habitat near Minne-
dosa, Manitoba, and the breeding population
increased from 12 pairs in 1969 to 66 pairs in
1971 on his 2.56-km2 study area. But only 9% of
the homing (marked, hand-reared) females at
Minnedosa produced broods in 1971. Sellers
attributed poor production to severe nest preda-
tion associated with a lack of nesting cover.
Upland nesting cover had been removed by
agricultural tilling and burning. Based on those
findings, the present study was initiated in 1971
on the Delta Marsh, where agricultural land use
is negligible and nesting cover appeared ade-
quate.
The objective of this study was to determine
whether hand-reared Mallards would home to a
specific release site in the Delta Marsh and, if so,
would reproduce at a level to maintain their
numbers. When it became evident in 1972 that
yearling hens were not producing broods, the
emphasis of the study was changed to a
comparison of productivity in hand-reared and
wild Mallards on the same areas.
Study Area
The Delta Marsh is a 140-km?2 expanse of
shallow bays, creeks, and potholes at the south
end of Lake Manitoba (Figure 1). The north
edge of the marsh is separated from the lake bya
sandy wooded ridge. Dense stands of Yellow
Cane (Phragmites australis) interspersed with
patches of White-top (Scolochloa festucacea)
meadow cover most of the dry surface from the
ridge to the bay edges. Cattail (Typha /atifolia)
grows in dense stands around potholes, creeks,
and on flooded bay shorelines. Bulrush (Scirpus
spp.) is common in large shallow bays. Addi-
tional description of topography, climate, and
vegetation is given by Bird (1961, p. 19), Love
and Love (1954), and Anderson and Jones
(1976).
In 1971 duckling releases were made at the
Diversion, 4.8 km W of the village of Delta. This
study area is 2.56 km? in size, divided by the
Assiniboine River Diversion. Creeks, potholes,
and borrow pits are typical wetlands on the area.
In 1972 an additional release site was estab-
56 THE CANADIAN FIELD-NATURALIST
ASSINIBOINE
Voles93
STUDY AREAS
| DIVERSION
2 AIKENS BAY
3 COOKS CREEK
@PAIRS OF YELLOW SADDLED MALLARDS
1 KM
+—4
FiGure |. Map of the Delta Marsh illustrating the study areas and the location of marked females established outside the
release sites in 1973.
lished at Aikens Bay, a 0.8-km? area, 8 km E of
Delta. The aquatic and upland habitat at Aikens
Bay was managed for waterfowl (mainly by
water-level regulation) and was considered to be
high quality. An unknown number of wild
Mallards occupied each release site prior to
stocking.
A 2.56-km? area at Cooks Creek Meadow,
4.8 km E of Delta, was chosen for a control (1.e.,
no birds released) in 1972. That area is typical
marsh habitat.
In 1973 water levels were lower than usual on
parts of the Delta Marsh. Wind tides from Lake
Manitoba maintained water levels in the larger
bays, but lack of precipitation left many nearby
potholes and shallow marshes dry. Precipitation
at Portage la Prairie was 7.62 cm lower than the
long-term mean for 6 mo prior to 30 March 1973
(Atmospheric Environment Monthly Reports).
Methods
Ducklings were hatched 1n an incubator, from
eggs taken from a captive flock originating from
eggs gathered in the wild and hatched at Delta.
Ducklings were hand-reared (Ward and Batt
1973) until 4-5 wk of age before release. Each
duckling was fitted with an individually num-
bered plastic nasal saddle and a standard U.S.
Fish and Wildlife Service leg band immediately
prior to release. Birds were liberated in groups of
10-15 throughout all wetlands on the study
areas.
In 1971, 456 females and 180 males were
released at the Diversion. In 1972, 503 females
and 34 males were liberated at Aikens Bay and
245 females at the Diversion site.
Mallard breeding densities were estimated by
checking on foot each wetland on the study area.
Each Mallard observed was checked for a nasal
saddle. Counts were conducted as suggested by
Dzubin (1969). A 12.87-km roadside transect
was run 5-6 times a week on the Diversion and
surrounding area each year and several walking
transects were conducted at Aikens Bay. These
transects provided information on the ratio of
marked to unmarked females, homing hens, and
the male:pair ratio. The ratio of marked to
unmarked females was used to obtain an
estimate of marked hens represented by drakes.
Sex ratio of Mallards counted prior to 15
1979
April was applied to lone males and groups of
five or less to correct breeding pair estimates for
the unmated drake cohort. I assumed that all
Mallards present were counted during each
census.
Several square kilometres of marsh sur-
rounding the study areas were searched each
year for marked birds. Locations of marked hens
were plotted on a map and revisited until the hen
was identified or had disappeared.
In 1973, nest searches were carried out 2-3
times a week between 09:00 and 12:00 from the
second week of April to the beginning of July.
Two observers and a Labrador retriever partici-
pated in each search. Nests were located, plotted
on a map, and marked witha piece of fluorescent
tape on vegetation or a stake 3-4 m away. Nest
initiation date was estimated for each nest by
back-dating eggs in incomplete clutches (Dane
1966). Eggs in nests containing down were
floated (Westerskov 1950) to estimate the ap-
proximate stage of’ incubation. Nests were
revisited after the anticipated hatching date and
nests containing finely crushed egg-shells and
membranes were considered hatched.
Brood searches consisted of walking the
emergents surrounding wetlands. The number
and age of young (in weeks) and the marking, if
any, on the female were used to avoid duplica-
tion in counts. This technique was supplemented
with morning and evening road transects,
walking transects, and observation periods
wherein one block of habitat was observed for
2-3 h. Broods in Aikens Bay were observed from
a 6-m tower in a central location on the site.
In spring 1973, 15 paired Mallard females
were collected on the marsh, at least 2 km away
from the study areas, in an attempt to estimate
the proportion of adults in the unmarked
population. Only females from isolated pairs
were shot. Hens were qualitatively classed as
yearling or adult based on examination of wing
feathers (Carney and Geis 1960; Hopper and
Funk 1970). Wings from known adults were
examined for comparison.
Results
Spring Arrival and Breeding Densities
The first Mallards (six) in 1972 arrived on the
Diversion on 7 April, and did not include any
BAILEY: WILD MALLARD STOCKING 57
TABLE I—Mallard breeding pair counts (with 95% con-
fidence intervals) on the Diversion and Aikens Bay, Delta,
Manitoba. Pairs are corrected for sex ratio.
Pairs
Study area 1971 1972 Total
released released unmarked
Diversion
5 May 1972 7 ae 7/ — 1947 36
17 May 1972 19+5 = 6} a8 5) ay
Means 18.0 13.5 32
1 May 1973 644 9+4 1145 26
17 May 1973 9+5 10+6 WD) ae 7/ 30
Means ES 9.5 LES 28
Aikens Bay
26 April 1973 —— 1444 il ae 5 35
8 May 1973 — 1344 9+5 22
Means 13.5 15.0 29
marked hens. On 13 April the first marked
female and her mate were observed in a flock of
five pairs on the study area. Most of the marked
Mallards did not arrive until the week starting 21
April, when the mean number of marked hens
observed per road transect increased to 3.85
from 0.58 the previous week. Over the same time
period the mean number of unmarked hens
observed per road transect decreased from 17.4
to 10.6.
Spring 1973 was phenologically very early,
and the first Mallards arrived at Delta on 21
March. On 27 Marcha marked yearling hen was
seen at Aikens Bay (Peter Ward, personal
communication). When I arrived on 12 April, all
hens were back.
The breeding pair censuses (Table 1!) were
timed to correspond with Mallard nest initiation
dates as indicated by increases in the male: pair
ratio along roadside transects. Prior to 15 April
1972, 496 Mallards were seen with a sex ratio of
57.3 males to 42.7 females. Prior to 15 April
1973, 543 Mallards showed a sex ratio of 54.7
males to 45.3 females.
Mallard breeding pairs on the Diversion
averaged 32 per 2.56 km? in 1972 and 28 in 1973
(Table 1). Aikens Bay contained 29 pairs of
Mallards (93 pairs per 2.56 km?2). Surveys
commencing 14 and 21 May 1973, on Cooks
Creek Meadow revealed a population of 54 and
55 pairs of Mallards per 2.56 km? respectively.
58 THE CANADIAN FIELD-NATURALIST
Homing of Mallards Released in 1971
and 1972
High pre-fledging mortality of an unknown
number of ducklings in 1971 severely limited the
number of hens alive to return in 1972 (Bailey,
unpublished data). In all, 28 females were
individually identified on the Diversion in 1972
(see below for number that actually settled on
the area). All homing females were accompanied
by unmarked drakes and no marked males were
observed. Extensive searches of the surrounding
marsh did not reveal additional marked birds.
Breeding pair counts indicated that about
eight 1971-released birds returned to the Diver-
sion in 1973, (Table 1). It was not possible to
recognize individually all 2-yr-old females in
1973 because the paint had worn off many
saddles. Assuming a 52% annual survival rate
for adult females in southwestern Manitoba
(Anderson 1975) and considering that 28 hens
returned to the Diversion in 1972, then the eight
birds observed in 1973 represented a homing rate
of 53% for 2-yr-old females. This adult homing
rate is considered minimal since the Delta Marsh
is a major Mallard harvest area within south-
western Manitoba.
In 1973, 17 hens released in 1972 were
identified on the Diversion (see below for
number that settled). Marked hens from 1972
releases there appeared to have homed solely to
the release site, because they were not found
elsewhere in the marsh.
Females homing to Aikens Bay encountered
high densities of marked and unmarked Mal-
lards (Table 1). At Aikens Bay, 14 returning
marked hens and | unmated marked male were
identified. An additional 17 marked hens were
observed scattered widely throughout the marsh
E of Delta and W of Clandeboye Bay (Figure 1).
Numerous marked females from Aikens Bay
releases only established themselves on the
lakeshore opposite Aikens Bay. Another
marked male was observed with an unmarked
hen at Marshy Point, 30.4km E of Delta
(Robert Blohm, personal communication).
I estimated that 186 of the females released at
the Diversion and 366 females in Aikens Bay
fledged in 1972. Robert Jones (personal com-
munication) calculated that the first-year mor-
tality rate of hand-reared Mallards released at
Delta between 1954 and 1970 was 67.2%. To
Vol. 93
calculate homing rate I assumed this mortality
rate for first-year fledged females in this study.
The 17 returning hens at the Diversion in 1973
represented 28% of the possible yearling sur-
vivors. There was a 26% homing rate (n = 31) of
marked yearling hens returning to Aikens Bay
and the east marsh in 1973.
Comparison of Breeding Pair Counts and
Homing Individuals
There was a discrepancy between the number
of identified yearlings homing to the Diversion
and the corresponding breeding pair estimates
(28 identified homing in 1972 vs. pair estimate of
18; and in 1973, 17 identified homing vs. pair
estimate of 9.5). This suggested that many
homing marked hens were missed during the
census or a certain proportion of marked
yearlings visited the study area only briefly each
year. In 1972, 13 (46%) of 28 homing hens were
identified only once during 51 roadside transects
on the Diversion whereas the remainder (15)
were recognized an average of 4.7 times each.
The mean residency bout for females sighted
more than once was 32 d in 1972, as determined
by the number of days between the first and last
sighting. In 1973, 7 (41%) of 17 hens were seen
only once during 53 transects and the remainder
(10) recognized an average of 2.5 times. The
mean residency bout was 31.3 d in 1973.
Nest Initiation and Productivity
Two of the earliest nests of unmarked hens
were back-dated to 16 and 20 April 1973,
whereas the earliest marked females (two) began
nesting on I5 May. A second peak of nest
initiation by unmarked hens occurred from 27
May to 2 June (Figure 2), and was apparently
due to renesting and initial nesting by Mallards
moving into newly-formed water areas after
3.2cm of precipitation received during this
period. A larger proportion of unmarked hens
than of marked yearlings had started nesting
prior to 15 May (x° = 4.8, P< 0.05, n = 36).
Generally poor reproductive success was
recorded for released birds nesting as yearlings.
Nest predators, chiefly Striped Skunks (Mephi-
tis mephitis), Raccoons (Procyon lotor), and
Red Foxes (Vulpes vulpes), were common on
both study areas and accounted for most nest
failures (Table 2). No marked Mallard hens with
broods were discovered on the Diversion in
1979
PERCENT OF TOTAL NESTS INITIATED
= I) Os £ i) o
Oo (e) Oo eo) eo) Oo
Oo
IS 2l
| APRIL | MAY
M= MARKED YEARLINGS n:=!!
Ml UNMARKED MALLARDS n= 24
MR
28% 9.7 12
BAILEY: WILD MALLARD STOCKING 59
Pi biba
19°26. 25° 79
| JUNE
I6 23
FIGURE 2. Distribution of nest initiation dates for marked vearlings and unmarked Mallards on the Delta Marsh study areas,
1973.
1972. In 1973, one brood of five ducklings was
found with a 1972-released hen; however, no
broods led by 1971-released hens were observed.
Two marked hens with broods were found at
Aikens Bay and two more marked hens with
broods were located in the vicinity of the release
site. One marked hen with a brood was observed
on Cooks Creek, 1.6 km S of Aikens Bay.
Mallard breeding pair counts for the
Diversion and Aikens Bay indicated that the
ratio of marked yearling hens to unmarked
females (41:40) was approximately I to I each
year. Significantly more broods accompanied by
unmarked hens (P = 0.005, binomial test, Siegel °
1956), however, were discovered on the release
sites.
TABLE 2—Nest success records for Mallards and other dabblers, Delta Marsh, 1973
Year Hatched Not
of
Species release Nests No. OK Destroved revisited! Deserted
Mallard
marked 1972 1] 2 20 8 | 0
marked 1971 3 0 0 3 - 0
unmarked 28 1] 42 IS) 2 0
Other
dabblers 81 23 36 32 19 7
'Not included in the calculation of the percentage of nests hatched.
60 THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 3—Percentage of marked and unmarked Mallard pairs producing broods at the Diversion and Aikens Bay in
1972 and 1973
1972 1973
Study area Year of Number of Percentage Number of Percentage
release breeding pairs producing broods breeding pairs producing broods
Diversion 1971 18.0 0.0 V3 0.0
1972 _— —_ QS 10.5
unmarked IBES 22. 15) 34.3
Aikens Bay 1972 _ — 13.5 14.8
unmarked — a 15.0 50.0
Fourteen nests of marked Mallards were Discussion
discovered during nest searches conducted in
1973 (Table 2). Eleven of these were nests of
yearling hens, and three were initiated by 2-yr-
old hens at the Diversion. Two of the 11 yearling
nests hatched, 8 were destroyed by predators and
1 was not relocated. All three nests of 2-yr-old
hens were destroyed.
Nesting success was 20% (n= 10) for nests
initiated by marked yearling Mallards, whereas
unmarked Mallard hens hatched 42% (n = 26) of
nests initiated (no significant difference
(P = 0.150) using a Fisher exact probability test
(Siegel 1956)). The low probability obtained,
however, suggests that the number of yearling
nests found may not constitute a large enough
sample to show a real difference, especially when
a substantial proportion (see below) of the
unmarked population was also yearling hens.
When the combined nesting success of I- and 2-
yr-old marked hens was compared with nesting
success of unmarked hens, the Fisher exact
probability decreased to 0.070. Nesting success
of other dabblers encountered during nest
searches was 36% (n = 81).
Fifteen paired Mallard females were shot in
spring 1973. These females included nine adults
and six yearlings, based on wing-feather exami-
nation. Evidently there was a large proportion of
adult females in the unmarked Mallard popula-
tion.
The percentage of unmarked hens producing
broods was much higher than that of marked
hens on each study area in both years (Table 3).
During the 2-yr study, 76 Mallard hens were
identified as first-year breeders; of these, only 6
hens (8%) were observed with broods on orinthe
vicinity of the release sites.
Most of the marked yearlings arrived well
after the majority of unmarked hens. Spring
arrival of hand-reared yearlings may extend over
several weeks. Sellers (1973) concluded that
yearling marked hens did not lag behind adults,
based on the appearance of marked birds with
the first arrivals in his area; however, he did not
determine when the majority of marked year-
lings arrived in relation to adults.
Wild-strain hand-reared Mallard females
were able to return to specific release sites within
the marsh. High pre-fledging mortality of 1971
releases did not prevent some survivors from
returning to the study area in spring 1972.
Homing of 53% (estimated) of surviving 2-yr-old
hens to the Diversion showed that nesting failure
during the previous year was probably not a
deterrent to homing in subsequent years. A high
percentage of adult hens homed in spite of low
water levels in 1973.
Lower homing rates of yearlings compared to
those of adults have been reported for Mallard,
Gadwall (A. strepera), Pintail (A. acuta), Blue-
winged Teal (A. discors) (Sowls 1955); Wood
Duck (Bellrose et al. 1964); Shoveler (A.
clypeata), (Poston 1974); and Tufted Duck (A.
fuligula) (Mihelsons et al. 1970). The pro-
portion of yearling marked Mallards homing
was approximately one-half that of marked
adults. Furthermore, only about 60% of homing
yearling hens resided on the area for extended
periods each nesting season. The remainder were
sighted only once, usually prior to nest initiation
or after nesting in July. These observations
support the contention of Hochbaum (1955, p.
124) that some hens may home to their natal area
but then move elsewhere to nest. On 29 July
1979
1973, a flightless, 1972-released Mallard was
observed 0.8km S of Aikens Bay, so an
unknown proportion of marked hens also used
the marsh to molt.
Males did not generally home to release sites.
In Mallards, pairing usually takes place on the
wintering grounds and males follow their mates
back to their natal area (Sowls 1955). But the
appearance of one homing drake on Aikens Bay
indicated that some drakes return to natal areas
if unmated in the spring.
Numbers of Mallards seen in May and June
aerial surveys of the marsh in 1973 increased 2.7
times over the previous three years’ average
(Delta Waterfowl Research Station, unpub-
lished data). This increase may have also been
partially due to low water levels causing birds to
be more visible than usual from the air. The
extremely high density of Mallard pairs at
Aikens Bay in 1973 was partly due to the homing
of release hens since one-half of the females
found there were marked. The unstocked Cooks
Creek Meadow also contained a relatively high
density of Mallard pairs. It is possible that
higher than usual numbers of Mallards on wet
areas of the marsh in 1973 masked the effects of
the stocking effort. In contrast, the Diversion
held comparatively few Mallards after two years
of stocking. The number of pairs on the
Diversion remained the same each year, and
there was evidence that marked yearlings were
homing to, but not remaining on, the site.
Marked yearlings also were found along the
lakeshore at Aikens Bay. It is likely that the
study areas were filled to capacity with breeding
pairs.
Reproductive behavior and success of first-
time breeders differs from those of adults in
many birds (Lack 1966). Bellrose et al. (1964)
found that yearling Wood Ducks nested later
than adults. Grice and Rogers (1965) showed
that yearling Wood Ducks were less successful
than adults in obtaining nesting sites when
breeding populations were high. Heusmann
(1975) indicated that limited nestings, smaller
clutch size, and lower brood survival led to
poorer annual production by yearling Wood
Ducks. Gates (1962) found that adult Gadwall
females established home ranges and began
nesting earlier than yearlings in the spring. Stotts
and Davis (1960) showed that adult Black Ducks
BAILEY: WILD MALLARD STOCKING 61
(Anas rubripes) were paired before yearlings and
suggested that adults reach breeding “tempo”
before hens nesting for the first time. Clutch size
was also larger in adult Black Ducks. Mihelsons
et al. (1970) found that yearling female Tufted
Ducks started nesting later than the older birds
and had comparatively poor success.
Yearling marked hens in this study arrived
later on the release sites than most unmarked
females, initiated nesting later, and were less
successful than unmarked hens in producing
broods. Lower productivity of released hens was
apparently not caused by a lack of nesting cover,
as found by Sellers (1973) because unmarked
hens on the same areas were much more
successful. Breeding by yearling hand-reared
Mallards resembles a pattern observed in first-
time breeders of other duck species, and lower
productivity may be partly due to the fact that
they are all breeding for the first time. The
assumption that a considerable proportion of
the unmarked Mallard population were adults
was supported by the collecting of 15 hens.
Poor nesting success was also documented for
a small sample of adult marked hens. Hence, the
possible effects of hand-rearing on future breed-
ing success remain unknown. It is important
from a management standpoint to determine
whether the differences observed between hand-
reared and wild Mallards in this study are due to
hand-rearing or age. The fact that no similar age-
related breeding biology studies of wild Mal-
lards are available to compare with these results
points out an important research need. In view
of the poor reproductive success of hand-reared -
birds and the apparently high potential for
natural immigration and production, Mallard
stocking 1s of questionable value on the Delta
Marsh.
Acknowledgments
I gratefully acknowledge the advice of J.R.
Bider received during this study. Thanks are also
due to B. D. J. Batt, P. Ward, and R. E. Jones
for suggestions during the field study, and to
them and A. J. Erskine, A. Dzubin, and R. D.
Titman for review of the manuscript. The help of
students and assistants at the Delta Waterfowl
Research Station was appreciated. The com-
ments and criticism of R.W. Stewart were
invaluable throughout this study. R. McCulloch
62 THE CANADIAN FIELD-NATURALIST
assisted with the illustrations. Support of the
Delta Waterfowl Research Station and Ducks
Unlimited Foundation is acknowledged. I grate-
fully acknowledge scholarships from the Na-
tional Research Council of Canada and the
Province of Québec.
Literature Cited
Anderson, D. R. 1975. Population ecology of the Mallard:
V. Temporal and geographic estimates of survival,
recovery and harvest rates. United States Fish and
Wildlife Service Resource Publication 125. 110 pp.
Anderson, M. G. and R. E. Jones. 1976. Submerged aqua-
tic vascular plants of the east Delta Marsh. Manitoba
Department of Renewable Resources and Transportation
Services. 120 pp.
Bellrose, F C., K. L. Johnson, and T. U. Meyers. 1964.
Relative value of natural cavities and nesting houses for
Wood Ducks. Journal of Wildlife Management 28:
661-676.
Benson, D. 1939. Survival studies of Mallards liberated in
New York State. Transactions of the North American
Wildlife Conference 4: 411-415.
Bird, R. D. 1961. Ecology of the aspen parkland of western
Canada in relation to land use. Canada Department of
Agriculture Contribution 27. 155 pp.
Carney, S. M. and A. D. Geis. 1960. Mallard age and sex
determination from wings. Journal of Wildlife Manage-
ment 24: 372-381.
Dane, C. W. 1966. Some aspects of breeding biology of the
Blue-winged Teal. Auk 83: 389-402.
Doty, H. A. and A. D. Kruse. 1972. Techniques for esta-
blishing local breeding populations of Wood Ducks.
Journal of Wildlife Management 36: 428-435.
Dzubin, A. 1969. Assessing breeding populations of ducks
by ground counts. /m Saskatoon wetland seminar.
Canadian Wildlife Service Report Series 6. pp. 178-230.
Errington, P.L. and W.E. Albert, Jr. 1936. Banding
studies of semi-domesticated Mallard ducks. Bird-
Banding 7: 69-73.
Foley, D. D. 1954. Studies on survival of three strains of
Mallard ducklings in New York State. New York Fish and
Game Journal |: 75-83.
Foley, D. D., D. Benson, L. W. DeGraff, and E. R. Holm.
1961. Waterfowl stocking in New York. New York Fish
and Game Journal 8: 37-48.
Gates, J. M. 1962. Breeding biology of the Gadwall in
northern Utah. Wilson Bulletin 74: 43-67.
Grice, D. and J. P. Rogers. 1965. The Wood Duck in
Massachusetts. Massachusetts Division of Fish and
Game, P-R Report Project W-19-R. 96 pp.
Vol. 93
Heusmann, H. W. 1975. Several aspects of the nesting
biology of vearling Wood Ducks. Journal of Wildlife
Management 39: 503-507.
Hochbaum, H. A. 1955. Travels and traditions of water-
fowl. University of Minnesota Press, Minneapolis. 301 pp.
Hopper, R. M. and H. D. Funk. 1970. Reliability of the
Mallard wing age-determination technique for field use.
Journal of Wildlife Management 34: 333-339.
Hunt, R.A., L.R. Jahn, R.C. Hopkins, and G.H.
Amelong. 1958. An evaluation of artificial Mallard pro-
pagation in Wisconsin. Wisconsin Conservation Depart-
ment, Technical Wildlife Bulletin 16. 70 pp.
Lack, D. 1966. Population studies of birds. Clarendon
Press, Oxford. 341 pp.
Lee, F.B. and A.D. Kruse. 1973. High survival and
homing rate of hand-reared wild strain Mallards. Journal
of Wildlife Management 37: 154-159.
Lincoln, F. C. 1934. Restocking of marshes with hand-
reared Mallards not proved practicable. United States
Department of Agriculture Yearbook 1934: 310-313.
Love, A. and D. Love. 1954. Vegetation of a prairie marsh.
Bulletin of the Torrey Botanical Club 81: 16-34.
Mihelsons, H., G Lejins, A. Mednis, and V. Klimpins.
1970. Attachment to the territory and reproduction
effectiveness of the Tufted Duck on Lake Engure (Latvian
SSR). International Congress of Game Biologists 30:
82-85.
Poston, H. J. 1974. Home range and breeding biology of
the Shoveler. Canadian Wildlife Service Report Series 25.
49 pp.
Schladweiler, J.C. and J. R. Tester. 1972. Survival and
behavior of hand-reared Mallards released in the wild.
Journal of Wildlife Management 36: 1118-1127.
Sellers, R.A. 1973. Mallard releases in understocked
prairie pothole habitat. Journal of Wildlife Management
37: 10-22.
Siegel, S. 1956. Nonparametric statistics for the behavioral
sciences. McGraw-Hill Book Company, New York. 312
Pp.
Sowls, L. K. 1955. Prairie ducks. Stackpole Company,
Harrisburg, Pennsylvania, and Wildlife Management
Institute, Washington, D.C. 193 pp.
Stotts, V. D. and D. E. Davis. 1960. The Black Duck inthe
Chesapeake Bay of Marvland: breeding behavior and
biology. Chesapeake Science I: 127-154.
Ward, P. and B.D. J. Batt. 1973. Propagation of captive
waterfowl. North American Wildlife Foundation and
Wildlife Management Institute, Washington, D.C. 64 pp.
Westerskoy, K. 1950. Methods tor determining the age of
game bird eggs. Journal of Wildlife Management 14:
56-67.
Received 12 June 1975
Accepted 2 August 1978
Notes
Recent Collections of the Black Redhorse, Moxostoma duquesnei,
from Ontario
EDWARD KotT,! ROBERT E. JENKINS,2 and GREGORY HUMPHREYS!
‘Department of Biology, Wilfrid Laurier University, Waterloo, Ontario N2L 3C5
2Department of Biology, Virginia Commonwealth University, Richmond, Virginia, USA 23284
Kott, Edward, Robert E. Jenkins, and Gregory Humphreys. 1979. Recent collections of the Black Redhorse, Moxostoma
duquesnei from Ontario. Canadian Field-Naturalist 93(1): 63-66.
The Black Redhorse, Moxostoma duquesnei, a species considered endangered in Canada, was recently collected from two
localities on the Nith River, a tributary of the Grand River, Ontario. It has not been collected in Canada since 1938. This
species is often confused with M. erythrurum, the Golden Redhorse. A comparison between these species is included.
Key Words: Moxostoma duquesnei, redhorse suckers, Ontario, morphology, endangered species.
The Black Redhorse, Moxostoma duquesnei, 1s ROM9637 Ontario, Elgin County, Catfish Creek. 26
generally found in clear streams of moderate width March 1937. H. C. White. 1 adult.
that are not subject to appreciable siltation. Because it © ROM10364 Ontario, Elgin County, Catfish Creek south
is susceptible to siltation, it is becoming rare in many of Aylmer. 6 March 1938. H.C. White. I adult or
E Underhill 1974; C d Colli supae ge
cone ee Sacske: 1969) Dea tes aa The Lake Champlain system of the St. Lawrence
. ; . drainage was included in the range of this species by
This species has previously been recorded in SES e
Canada from only two localities in Ontario, both in Hubbs (1930; p. 23) onthe basis of “Greeley (MIS) Its
the Lake Erie drainage: a tributary of the Grand River poppe ae musucalice! by Uaioe (ER BHT Cre) WO
(Hubbs and Brown 1929) and from Catfish Creek miclnge mul OH UNS Sie Lanitenge re RV: Gresley
(Hubbs and Brown 1929; Scott and Crossman 1973). CHG ioe (pHIOLS SWieln ers! aun ee
The last recorded capture of this species in Canada SOND fia VINE Lorene dramage) Woearmutcd
was in 1938 and repeated attempts to collect it from ee tate ele Crain eheavine ise (On) the Conese:
Catfish Creek since then have failed (Scott and oe a ee le ae ne,
Crossman 1973). For this reason the species has been peas ) als@ lundhealios (ie MeN ete So
included in lists of rare and endangered Canadian Grune Wes not neal peste) Byjthespeoe
fishes (Anonymous 1970: McAllister and Gruchy Three specimens of this species were recently
1977). horn La eren collected by two of us from two localities in the Nith
ys eae: Beene ie rae one. River, a tributary of the Grand River, in the Lake Erie
McAllister and Gruchy 1977). drainage basin, at least 35 km upstream from the »
Specific locality records of previous collections of sae ey 28) soe ne ue ee pace
this species based on specimens in the University of SOaTe So olantorn, coe) lana Sete ue! Melati (Cg
Michigan, Museum of Zoology (UMMZ) and Royal 1), was collected on 21 October 1976 ESE woWmn ee
Ontario Muweun GROMD ake as Holla: Plattsville, Oxford County, Ontario, (43 18 30 N;
: 80°37'30”W) using a 9-m, 6.4-mm-mesh seine. This
UMMZ85887_ Ontario, Catfish Creek, | mi W of Jaffa. part of the river consists of a stretch of swift-flowing
21 July 1927. Brown and Rupp. Total 45 spms., 1 ad — shallow water (about 15 cm deep at the time) over a
(202 mm SL) others young (reported by Hubbs and gravel bottom. This section ends in a poo] about | m
Brown 1929), as 56 specimens. deep with a rubble bottom. The specimen, which was
UMMZ89075 Ontario, Oxford County (near Brant Coun- —_—co}lected in the shallows, is in the Wilfrid Laurier
ty line) Cedar Creek, tributary of Grand River between University Museum as number WLU5594.
Paris and Princeton. 5 Sept. 1928. C. L. Hubbs. 3 speci- Two specimens (81mm and 79 mm_ standard
mens 75, 215, 215 mm SL (3 Aug. 1927 according to : : 5
length) were collected using the same technique on 29
Hubbs and Brown 1929) [no stream in the area indicated ; Sipe
currently bears the eae Cheeta June 1977 (Figure 1) at the second locality just north
ROMI975 Ontario, Elgin County, Catfish Creek near Of Ayr, Waterloo County, (43° 17'45"N; 80°28°10°N)
Aylmer. 21 March 1926. H. C. White. 1 specimen, 253. and 24 km downstream from the first. This segment of
SL. the river is similar to the previous in that it consists of
63
64 THE CANADIAN FIELD-NATURALIST
) ith il
cm ]
AIL
a | sl | al ie
Vol. 93
|
FIGURE 1. Moxostoma duquesnei collected from the Nith River: upper specimen, NMC 78-1; middle, WLU5233: lower,
WLU5594.
a zone of rapids followed by a deep pool. The rapids
had a boulder bottom with a depth up to | m. At
low water levels, the pool was effectively divided bya
gravel bar into two smaller pools. At the time of
capture the depth of the pool was about 75 cm. The
sides were gravel and the bottom was covered in fine
silt. Both specimens were collected from the pool. One
specimen is deposited in the Wilfrid Laurier Uni-
versity Museum as WLU5233, and the other at the
National Museum of Natural Sciences, Ottawa as
NMC78-1.
Moxostoma erythrurum, Golden Redhorse, was
collected with M. duquesnei at the first locality and
M. macrolepidotum, Shorthead Redhorse, was col-
lected at the second locality.
The earlier and recently collected specimens of M.
duquesnei agree in morphology and coloration with
the species from elsewhere in the Lake Erie basin
(tributaries in the United States) and from through-
out most of the wide range of the species in the United
States, from the southern parts of the lakes Ontario,
Huron, and Michigan basins and southward.
The species in the Nith River system that most
closely resembles M. duquesnei is M. erythrurum.
Many workers encounter difficulty in distinguishing
them. The following meristic data (taken by methods
of Hubbs and Lagler 1958) are from specimens from
the Lake Erie basin of Canada and the United States
unless otherwise stated (Jenkins 1970 and subsequent
study).
The best character for separation of these two
species is the lateral-line scale count: M. duquesnei
ranges 45-48, x = 46.4, N = 28: M. eryvthrurum 39-44,
x = 41.0, N = 46. Slightly overlapping counts may be
expected from additional specimens. Other meristic
differences are the following: body circumferential
scales (axis just anterior to dorsal fin), M. duquesnei
33-36, M. erythrurum 29-34; predorsal scale rows, M.
duquesnei 16-20, M. erythrurum 13-16; pelvic fin
rays, M. duquesnei usually one or both fins with 10
rays (in 19 of 28 specimens; nine with 9 as the highest
count), M. erythrurum 9-9 or 9-8 in 50 of 52
specimens, 10-9 or 10-10 in the other two; post-
Weberian vertebrae (counts from throughout total
range of species), M@. duquesnei 39-43 (40 in largest
Nith specimen), M. erythrurum 35-39.
1979
NOTES 65
TABLE 1—Some characteristics of three specimens of Moxostoma duquesnei and 11 specimens of M. ervthrurum from the
Nith River
Specimen Standard. Head length/ Eye diameter/ Caudal peduncle Caudal peduncle Number of Number of
and museum length, snout length snout length depth/ depth/ lateral pelvic rays
number mm caudal peduncle standard length line Slctimnrichit
length scales ia
M. duquesnei
WLU5594 49 2.84 0.81 0.6 0.094 47 10 10
WLU5233 79 2.56 0.55 0.67 0.090 46 9 9
NMC78-1 81 Dee) 0.61 0.7 0.095 46 9 9
Mean 69.7 2.65 0.66 0.66 0.093 46.3 OS OS
Range 49-81 2.55-2.84 0.55-0.81 0.6-0.7 0.090-0.095 46-47 9-10 9-10
M. erythrurum
WLU4763 41 3.3 0.94 0.68 0.102 40 9 9
45 3323 0.94 0.67 0.104 42 9 9
47 3.29 0.94 0.68 0.094 42 9 9
49 3.18 0.85 0.69 0.100 43 9 9
49 323 0.89 0.68 0.090 43 9 9
WLU4556 54 2.62 0.69 0.72 0.106 44 9 9
57 Des 0.65 0.66 0.093 42 10 10
59 2.90 0.76 0.71 0.102 43 9 9
62 2-38) 0.64 0.71 0.103 42 10 9
62 2.62 0.68 0.68 0.094 4] 9 9
WLUS5305 75 3.02 0.87 0.91 0.111 40) 9 9
Mean 54.6 2.92 0.80 0.71 0.100 41.8 O22 ON
Range 41-75 2.53-3.3 0.64-0.94 0.66-0.91 0.090-0.111 40-44 9-10 9-10
The species also differ in certain aspects of
morphometry (Hubbs and Lagler 1958; Jenkins 1970;
Phillips and Underhill 1971; Scott and Crossman
1973; Pflieger 1975) but greater overlap occurs with
these characters than in at least most of the former.
Some of the overlap is due to allometric growth of
body parts and to comparison of samples of speci-
mens with dissimilar body lengths (Jenkins 1970).
Table | expresses the interspecific differences, with
overlapping, based on the young and small juveniles
from the Nith River.
A recently discovered difference, that is particularly
useful for sorting large series of small specimens,
concerns coloration of the upper body. The difference
is best developed in the anterior dorsolateral area. In
M. duquesnei the melanophores of each scale are
more uniformly distributed, the total effect being an
evenly dusky to darkly colored back. In M. ery-
thrurum the posterior margin of each scale usually is
obviously darker than the somewhat pale central area
of the exposed portion of the scales, hence the scales
appear somewhat bicolored. The difference is usually
slight, but is discernible using syntopically collected
and similar-sized specimens of the two species.
The size of the specimens collected, apparently
young and small juveniles, indicates that a breeding
population of this species still exists in Canada.
The Nith River follows a series of gravelly spillways
until it unites with the Grand River (Chapman and
Putnam 1966). For this reason it has a considerable
gradient, with a minimum summer flow of about
2.8 m>s!. The region drained is rural in nature, with
New Hamburg (population 2990 in 1974) being the
largest center.
For part of its course it follows valleys 15.2-30.5 m
in depth: the other stretches are bounded by flood-
plains. These floodplains are largely cleared and are
used for growing grain or as pasture land; however, in
some areas natural reforestation 1s occurring on
abandoned or little-used farmland.
The river system is relatively clear, except during .
the spring flood period. Turbidity in Formazin
Turbidity Units, for June and July, taken at a location
near Ayr collection site was 2.70 and 2.40 (Anony-
mous, no date).
The Nith River system should be protected to
enable the continued existence, in Canada, of the
ecosystem that has harbored this species and other
more typical elements of the biota of Canada. This
species, itself, should be protected as an endangered
species in Canada.
Acknowledgments
We thank Frank Mallory, Department of Biology,
Wilfrid Laurier University and Bruce M. Buckland,
Ministry of Natural Resources, Cambridge, for
assisting us in collecting the specimens. Additional
specimens were kindly loaned by E. J. Crossman,
66 THE CANADIAN FIELD-NATURALIST
Royal Ontario Museum, Toronto, and R. M. Bailey,
University of Michigan, Museum of Zoology, Ann
Arbor. D. E. McAllister provided the X-ray negative
used for the vertebral count. This study was sup-
ported, in part, by National Research Council of
Canada Grant No. A-0575 to E. Kott.
Literature Cited
Anonymous. Undated. Water quality data for Ontario
lakes and streams. 1974. Volume IX. Water Resources
Branch, Ontario Ministry of the Environment. xxxiii +
324 pp.
Anonymous. 1970. Table of extinct, rare and endangered
vertebrates in Ontario. Ontario Fish and Wildlife Review
9(1-2): 15-21.
Chapman, L. J. and D. F. Putman. 1966. The physiog-
raphy of southern Ontario. Second edition. University of
Toronto Press, Toronto. xiv + 386 pp.
Cross, F.B. and J.T. Collins. 1975. Fishes in Kansas.
University of Kansas Museum of Natural History, Public
Education Series Number 3. 189 pp.
Eddy, S. and J.C. Underhill. 1974. Northern fishes.
University of Minnesota Press, Minneapolis. 414 pp.
Harlan, J. R. and E.B. Speaker. 1969. lowa fish and
fishing. State Conservation Commission, Des Moines,
lowa. 365 pp.
Hubbs, C. L. 1930. Materials for a revision of the cato-
stomid fishes of eastern North America. University of
Vol. 93
Michigan Museum of Zoology Miscellaneous Publica-
tion 20. 47 pp.
Hubbs, C. L. and D. E.S. Brown. 1929. Materials for a
distributional study of Ontario fishes. Transactions of the
Royal Canadian Institute 17. 56 pp.
Hubbs, C.L. and K.F. Lagler. 1958. Fishes of the
Great Lakes region. University of Michigan Press, Ann
Arbor, Michigan. 213 pp.
Jenkins, R. E. 1970. Systematic studies of the catostomid
fish tribe Moxostomatini. Ph.D. thesis, Cornell
University, Ithaca, New York. 818 pp.
McAllister, D.E. and C.G. Gruchy. 1977. Status and
habitat of Canadian fishes in 1976. Jn Canada’s threatened
species and habitats. Canadian Nature Federation,
Special Publication 6. pp. 151-157.
Pflieger, W.L. 1975. The fishes of Missouri. Missouri
Department of Conservation. 343 pp.
Phillips, G. L. and J. C. Underhill. 1971. Distribution and
variation of the Catostomidae in Minnesota. Bell Museum
of Natural History, University of Minnesota, Occasional
Papers, Number 10: 1-45.
Scott, W. B. and E. J. Crossman. 1973. Freshwater fishes
of Canada. Fisheries Research Board of Canada Bulletin
184. 966 pp.
Trautman, M. B. 1957. The fishes of Ohio with illustrated
kevs. Ohio State University Press, Columbus, Ohio. 683
PP-
Received 8 June 1978.
Accepted 8 August 1978.
Xanthochroism! in the Evening Grosbeak
CHRISTOPHER W. HELLEINER
Nova Scotia Bird Society, c/o Nova Scotia Museum, 1747 Summer Street, Halifax, Nova Scotia B3H 3A6
Helleiner, Christopher W. 1979. Xanthochroism in the Evening Grosbeak. Canadian Field-Naturalist 93(1): 66-67.
Key Words: xanthochroism, Evening Grosbeak (Hesperiphona vespertina).
This note describes two unusually pigmented
Evening Grosbeaks, Hesperiphona vespertina, and
summarizes several other reports between 1970 and
1977 in Nova Scotia. Xanthochroism in this species
has previously been reported by Saunders (1958) and
Godfrey (1967).
L.B. Macpherson (Maitland, Hants County, Nova
‘According to Harrison (Bird-Banding 1966, 37: 121) the use
of the term “xanthochroic” should be limited to replace-
ment of normal color by yellow. Most examples reported
are more probably non-melanic schizochroic, where the
absence of melanin leaves carotenoid pigments pheno-
typically apparent. I have therefore inserted the word
“apparent” to modify xanthochroism in the text.
Scotia) described an almost completely lemon-yellow
and white male Evening Grosbeak in winter 1969-70.
The primaries and tail were entirely white, except for
the tip of one tail-feather, which was dark brown or
black. The head was faintly tinged with olive, but the
rest of the plumage was yellow. The legs and feet as
well as the lower mandible were flesh-colored. The
upper mandible was gray. The eyes were dark. The
same bird, or one very like it, reappeared the following
winter.
I saw a xanthochroic male Evening Grosbeak with
somewhat more dark pigmentation in Halifax for a
few days on and around 19 March 1977. The dark on
the head, breast, and back were largely absent, except
for a little coarse brown mottling on a yellow
ISS)
background. The dark color in the wings and tail
appeared normal, but the white area seemed some-
what more extensive than usual, overlapping the dark
primaries a little.
Both the Maitland and the Halifax bird behaved
normally in the company of others of their species,
and competed aggressively for food.
Less detailed reports of several other apparently
xanthochroic Evening Grosbeaks have appeared in
winter bird reports in the Nova Scotia Bird Society
Newsletter in 1972, 1973, and 1977. These represent a
total of about seven such individuals since the first
specimen was collected in this province in 1958.
The Evening Grosbeak has become much commoner
in Nova Scotia during the last two decades. Average
numbers recorded on Christmas counts have in-
creased from 330 to 1160 to 1610 per count in
successive 5-yr periods starting in 1963 (Nova Scotia
Bird Society Newsletter). One might therefore expect
aberrant individuals to be seen more often. Neverthe-
less the frequency with which this otherwise very rare
NOTES 67
color variation is being reported in this species is
surprising. Smith (1966) states that xanthochroism is
rarer than other color variations in wild birds; Gross
(1965) lists many examples of melanism, several of
erythrism, but only six of xanthochroism. Perhaps in
the Evening Grosbeak affected individuals do not
suffer significant competitive disadvantage, and live
to reproduce.
Literature Cited
Godfrey, W.E. 1967 Xanthochroism in the Cape May
Warbler and Evening Grosbeak. Canadian Field-Natural-
ist 81: 226-227.
Gross, A. O. 1965. Melanism in North American birds.
Bird-Banding 36: 240-242.
Saunders, A. A. 1958. A yellow mutant of the Evening
Grosbeak. Auk 75: 101.
Smith, R. W. 1966. A case of xanthochroism in the Yellow-
faced Grassquit, Tiaris olivacea. Ibis 108: 627-628.
Received 19 May 1978
Accepted 18 September 1978
Interaction between a Long-tailed Weasel and a Snowy Owl
PETER C. BOXALL
Department of Biology, University of Calgary, Calgary, Alberta T2N IN4
Boxall, Peter C. 1979. Interaction between a Long-tailed Weasel and a Snowy Owl. Canadian Field-Naturalist
93(1): 67-68.
A Long-tailed Weasel (Mustela frenata) was observed rushing towards a Snowy Owl (Nyctea scandiaca) and its recently-
captured prey. This interaction is interpreted as attempted food piracy. The fearlessness of weasels may account for their
presence in the diet of owls.
Key Words: Mustela frenata, Nyctea scandiaca, food piracy, predator.
Little is known about the relationships of weasels
(Mustela spp.) to other predators. But the aggressive
predatory behavior and curiosity of weasels have been
described (Armitage 1961; Hansen 1952). While
conducting research on the behavioral ecology of
wintering Snowy Owls (Nyctea scandiaca), | wit-
nessed an interaction between a weasel and a Snowy
Owl.
At 11:24 on 15 December 1976, I was observing a
large female Snowy Owl perched on a fence-post at
the edge of a stubblefield, 6 km E of Calgary, Alberta.
I noted a Long-tailed Weasel (M. frenata) moving
slowly through the field approximately 30 m from the
owl. At 11:25 the owl flew into the stubblefield,
captured a small mammal, and remained on the
ground with the prey item in its feet. During this time,
the weasel was sitting upright about 20 m from the
capture location, and had apparently witnessed the
hunting attempt. After the capture, the weasel slowly
moved to within 10m of the owl. As the weasel
approached, the owl picked up the prey in its bill,
shook it several times, and started to swallow it. The
weasel then ran quickly to within 3 m of the owl,
whereupon the owl flew back to its original perch. The
weasel continued forward and investigated the area
where the hunting attempt occurred. Subsequently,
the owl flew 500-600 m away and perched in a large
tree.
Both owls and weasels are dependent upon rodents
for food during the winter. Weasels, however, are
68 THE CANADIAN FIELD-NATURALIST
sometimes utilized as food by wintering Snowy Owls.
Gross (1944) reports the remains of three weasels
(species not given) in one Snowy Owl stomach.
Catling (1973) found the stomach of one Snowy Owl
in Ontario to contain an entire Ermine (M. erminea).
In addition, | have found the remains of one Least
Weasel (M. nivalis) and three Long-tailed Weasels in
a sample of 100 pellets from the Calgary region.
Although the vision of weasels appears to be quite
poor (Murie 1935), their ability to locate moving
objects, and to detect prey by scent, is good (Ewer
1973). The weasel, during the interaction described
above, behaved as if the owl’s prey was detected by
scent. I believe that the weasel’s actions towards the
owl were probably motivated by hunger.
The sudden rush by the weasel at the owl as the
latter started swallowing the prey suggests a possible
attempt at food piracy. The unusual behavior of the
weasel, and the fact that the owl had just ingested a
prey item, probably allowed the weasel to escape
unscathed. Fearless behavior toward a larger pred-
ator, as described above, could explain in part the
observed levels of predation upon weasels in nature.
Vol. 93
I thank M. Ross Lein, Dick Braithwaite, and Tom
Dickinson for criticism of the manuscript. This
observation was made while I was conducting
research supported in part by grants from the
National Research Council of Canada to M. Ross
Bem:
Literature Cited
Armitage, K. B. 1961. Curiosity behavior in some muste-
lids. Journal of Mammalogy 42: 276-277.
Catling, P.M. 1973. Food of the Snowy Owl wintering in
southern Ontario, with particular reference to the Snowy
Owl hazard to aircraft. Ontario Field Biologist 27: 41-45.
Ewer, R.F. 1973. The carnivores. Cornell University Press,
Ithaca, New York.
Gross, A.O. 1944. Food of the Snowy Owl. Auk 61:
1-18.
Hansen, R.M. 1952. Notes on behavior of long-tailed
weasel in Utah. Journal of Mammalogy 33: 492.
Murie, A. 1935. A weasel goes hungry. Journal of Mam-
malogy 16: 321-322.
Received 23 May 1978
Accepted 3 August 1978
Black-necked Stilts Nesting near Edmonton, Alberta
Dick DEKKER,! ROBERT LISTER,2 TERRY W. THORMIN,*.D. V. WESELOH,? and LINDA M. WESELOH4
13819-112A St., Edmonton, Alberta T6J 1K4
210823-80 Avenue, Edmonton, Alberta T6E 1V9
3John Janson Nature Centre, Edmonton Parks and Recreation, Edmonton, Alberta. Present address: Provincial Museum of
Alberta, Edmonton, Alberta TSN 0M6
4Provincial Museum of Alberta, Edmonton, Alberta T5N 0M6. Present address: Canadian Wildlife Service, Canada Centre
for Inland Waters, P.O. Box 5050, Burlington, Ontario L7R 4A6
Dekker, Dick, Robert Lister, Terry W. Thormin, D. V. Weseloh, and Linda M. Weseloh. 1979. Black-necked Stilts nesting
near Edmonton, Alberta. Canadian Field-Naturalist 93(1): 68-69.
A series of sightings of Black-necked Stilts, Himantopus mexicanus, near Edmonton, Alberta, in the spring of 1977
culminated in the finding of two nests with eggs, which provide the first authenticated records of this species breeding in
Canada.
Key Words: Himantopus mexicanus, breeding record, Black-necked Stilts, Alberta.
On 1 May 1977, when checking birds with
Kathleen Ball and Bruce and Karen Heming at the
southeast end of Beaverhill Lake some 80 km (50 mi)
SE of Edmonton, RL was told by Ron Slagter of
Edmonton that he had found two Black-necked Stilts
on a shallow bay of the lake. They proceeded to the
spot and saw the birds feeding with about 20
American Avocets, Recurvirostra americana. The
stilts were viewed for about 20 min with binoculars
and telescope at a distance of 50 m, but the birds were
readily identified with the naked eve. Later it was
learned that the birds had also been seen by Gerry and
Betty Bulmer of Edmonton, who took recognizable
photographs of them (Bulmer 1977).
On 3 May, RL was informed that R. C. Hodgins, of
Stony Plain, Alberta, had seen another pair of stilts
ona pond near St. Albert, about 10 mi(16 km) NW of
Edmonton, but a later search of the area failed to
relocate the birds. Hodgins is familiar with the stilt
and was positive of his identification.
1979
On 24 May, Dick Dekker reported two Black-
necked Stilts on the north shore of Beaverhill Lake,
and on 4 June, he found four of the birds in a marshy
area on the west side of the lake. On the following day,
Dekker and Lister went to view the birds. They were
joined at the lake by the Weselohs and Thormin.
The area had been dry in April, but heavy rains and
some snow later had flooded it, so in June it was mostly
mud and water, in parts heavily overgrown with
Marsh Ragwort (Senecio congestus var. palustris). At
the marsh were three Black-necked Stilts and several
avocets. One of the stilts seemed to run from a clump
of ragwort, and resorted to a “broken-wing” dis-
traction display. We withdrew for a time to give the
bird a chance to return to a nest if it had one. After
about 20 min we moved in, and again the bird was seen
to run from a mound of mud. DVW waded through
the mire and was rewarded by finding a nest and four
eggs. He collected one egg which is now in the
Provincial Museum of Alberta, and pictures were
taken of the nest.
The nest was made of weed stems, apparently the
stalks of ragwort, and was located just above the water
level. Avocets’ nests in the vicinity were all on dry land
some distance from the water, which may have
receded since the eggs were laid. All the avocet nests
we saw had less nest material in them than that of the
stilt.
On 17 June DVW and Eric Tull discovered a second
stilt nest approximately 50 m west of the first one. The
second nest contained seven eggs, one of which was
also collected and preserved at the museum. This nest,
except for the greater number of eggs, was similar in
construction, content, and placement to the first.
Three days later the first nest contained only one egg,
which was pipped. By 07:45 on 21 June the chick had
successfully emerged and was observed in the nest
(DVW and Eric Mills). The fate of the young from the
other two eggs is unknown, but they probably hatched
on 19 or 20 June and were led to the nearby marsh. By
23 June the first nest was completely vacated and the
second nest still contained six eggs. Hatching in the
second nest was first noted at 06:10 on 4 July when the
nest contained one young and five eggs. By 20:15 on 4
July, two young had emerged and a third could be
heard peeping inside its cracked egg; the two young
were banded by Martin McNicholl. On 7 July the
second nest still contained three eggs which were cold
and addled, and the remains of a unbanded young stilt
were found only a few feet from the nest. Presumably
this was the young that was heard peeping on 4 July.
The three addled eggs were collected and are
preserved at the museum.
The two nests of Black-necked Stilts near Edmon-
ton, in 1977 contained I 1 eggs, of which four definitely
hatched, two probably hatched, two were collected
NOTES 69
early in incubation, and three were addled (and
subsequently collected). Three young are known to
have left the nest, two others may have, and one was
found dead.
In the spring of 1977 there were far more avocets
around Beaverhill Lake than RL could remember in
about 50 years of visiting the lake (see also Dekker
1977). Dry weather in the western states as well as in
sourthern Alberta, accompanied by very little run-off
from the meagre snowfall of winter, had left many
marshes dry, and it is possible birds had travelled far
north of their normal breeding grounds to find
suitable nesting habitat. There were also several
sightings of Black-necked Stilts in Montana during
the first week of May, and that state too recorded its
first breeding records (two) of the stilt in 1977 (Paul
Skaar, personal communication to DVW).
Salt and Salt (1977) list three previous sightings of
the Black-necked Stilt in Alberta. Parts of a Black-
necked Stilt are said to have been found (by Tom
Randall) near Brooks in the mid-1950s but they were
not preserved. On 12 May 1970, a bird was seen and
photographed near Calgary (Weseloh 1972) and on 24
May 1972, another was seen and photographed near
Irricana (Weseloh and McKay 1972).
Godfrey (1966) remarks “A set of eggs in the
National Museum of Canada was, according to the
label, taken at Qu’Appelle, Saskatchewan, on June
13, 1894, by Edward Arnold. This is so far from the
known breeding range that the possibility of an error
in the data seems probable.” In light of the current
Alberta and Montana nesting records, perhaps the
Saskatchewan record should be given more credence.
Literature Cited
Bulmer, B. 1977. Black-necked Stilts observed at Beaverhill
Lake. Edmonton Naturalist 5(6): 145.
Dekker, D. 1977. Avocets and habitat. Edmonton
Naturalist 5(6): 144.
Godfrey, W.E. 1966. The birds of Canada. National .
Museum of Canada Bulletin 203. 428 pp.
Salt, W.R. and J. R. Salt. 1977. The birds of Alberta.
Hurtig Publishers, Edmonton.
Weseloh, D. V. 1972. First verified record of the Black-
necked Stilt for Alberta. Canadian Field-Naturalist
86:165.
Weseloh, D. V. and W.G. McKay. 1972. The second
report of a Black-necked Stilt in Alberta. Calgary Field-
Naturalist 4(5):; 90-91.
Received 10 April 1978
Accepted 6 August 1978
Addendum
No Black-necked Stilts were seen at Beaverhill Lake
in 1978.
Received 8 November 1978
70 THE CANADIAN FIELD-NATURALIST
Vol. 93
Common Garter Snake Predation on Robin Nestlings
KATHY MARTIN
Department of Zoology, University of Alberta, Edmonton, Alberta
Present Address: Biology Department, University of Prince Edward Island, Charlottetown, Prince Edward Island CIA 4P3
Martin, Kathy. 1979. Common Garter Snake predation on Robin nestlings. Canadian Field-Naturalist 93(1): 70-71.
Key Words: American Robin, Turdus migratorius, nests, nestlings, Common Garter Snake, Thamnophis sirtalis, predation,
prey selection.
During a two-year study of American Robins
(Turdus migratorius) on Comox burn, a Douglas Fir
(Pseudotsuga menziesii) plantation on Vancouver
Island, British Columbia, a large percentage of nests
were unsuccessful. Of 175 active nests found during
1971 and 1972, at least 59 (34%) were destroyed owing
to predation (Martin 1973). The only predation I
actually witnessed involved a Common Garter Snake
(Thamnophis sirtalis) removing young Robins froma
nest on the ground.
On 11 July 1971 at 16:15 I found a Robin nest well
camouflaged under a fallen log. Three nestlings
approximately 9 d old begged vigorously when
approached, and I banded, weighed, and measured
them. Their weights were 32, 35, and 54 g. On 12 July
at 09:40 I found that a garter snake, approximately
90 cm long, had removed the smallest nestling and
was ingesting it. Another young was out of the nest
and the third remained in it. As I approached, the
snake began to move away. It had ingested the
nestling’s head and neck, but the abdomen and legs
were still dangling from its mouth. I wished to detain
the snake for Martin McNicholl (MKM) to photo-
graph, so I tied a string to the nestling’s leg, attached it
to a Red Huckleberry (Vaccinium parvifolium) stem
and left the site. At 10:05 the snake had regurgitated
the Robin and disappeared. I returned the live young
to the nest, collected the dead one, and entered a small
observation blind. At 11:30, I heard a nestling
squawking and saw a snake pull the second lightest
young (35 g) from the nest. The snake was biting or
grasping the neck of the nestling as MKM and I
approached. When the snake saw us, it tried to move
away with the young Robin, but the nestling
prevented this by grabbing vegetation with its feet.
MKM took several photographs and we left the nest
site. As I watched from a blind, the snake pulled its
prey a short distance away and out of sight. At 12:00
the female Robin returned with food in her beak and
remained near the nest chirping softly until 12:20
when she fed the remaining young (54 g) and flew
away. The snake crawled out of a hole near the nest at
12:30 at which time it had ingested only the head and
neck of the second nestling. At 12:50, the male Robin
returned with food, but by then the snake had
disappeared. At 13:45 I revisited the site and found the
second nestling dead 6 m north of the nest. As there
was no interference by humans or adult Robins after
11:30 it may be that the snake was unable to swallow
this nestling. At 15:35, the third and largest young was
still in the nest, and on 13 July at 11:10 found it fledged
about | m north of the empty nest. At 12:00 on 13
July, the second dead young Robin was still near the
nest and I saw a snake nearby of the same species and
size as seen earlier. On 15 July at 15:45, the dead
nestling was gone.
While I witnessed the predation, the adult Robins
did not observe the snake, because they spent
relatively long periods away from the nest and the
snake was not present when they returned. The
literature refers to snakes as occasional predators of
Robin eggs and young (Bent 1949). On my study area,
predation on Robin nests by garter snakes was
probably infrequent because most Robin nests were
situated approxmately | m above the ground in trees
(Martin 1973). The major prey species of T. sirtalis on
Vancouver Island are amphibians and earth worms
(P. Gregory, personal communication): however, T.
sirtalis does occasionally eat nestling birds, sometimes
climbing small bushes and trees (Gregory 1975).
There have been a number of reports of snakes
raiding passerine nests (Cink 1977: Best 1974, 1978:
Ervin and Rose 1973). Both Cink (1977) and Best
(1974) reported that snakes ate the complete clutch of
eggs or young. Best (1974) reported Blue Racers
(Coluber constrictor) returning to the nest after they
had removed all the contents. I have found no reports
that snakes have shown selectively in killing nestlings.
My observations indicated that a snake first killed the
lightest and smallest nestling. It then returned and
killed the second-lightest nestling, possibly indicating
prey selection by size. Because the snake abandoned
the second nestling, perhaps these young Robins were
larger than normal prey and too large to ingest.
I thank P.T. Gregory, M. Harris, M. K. Mc-
Nicholl, and F.C. Zwickel for comments on the
manuscript.
1979
Literature Cited
Bent, A. C. 1949. Life histories of North American thrush-
es, kinglets, and their allies. United States National
Museum Bulletin Number 196. 452 pp. (1964 Dover
reprint consulted.)
Best, L. B. 1974. Blue Racers prey on Field Sparrow nests.
Auk 91: 168-169.
Best, L. B. 1978. Field Sparrow reproductive success and
nesting ecology. Auk 95: 9-22.
Cink, C. L. 1977. Snake predation on Bell’s Vireo nest-
lings. Wilson Bulletin 89: 349-350.
Ervin, S. and C. Rose. 1973. Gopher Snake predation on
NOTES ial
the Common Bushtit. Auk 90: 682-683.
Gregory, P. T. 1975. Arboreal mating behavior in the Red-
sided Garter Snake. Canadian Field-Naturalist 89: 461-
462.
Martin, K. 1973. Breeding density and reproductive success
of robins in relation to habitat structure on logged areas
of Vancouver Island, British Columbia. M.Sc. thesis,
University of Alberta, Edmonton. 89 pp.
Received 19 May 1978
Accepted II] August 1978
Distribution and Movements of Selkirk Caribou, 1972-1974
DAVID J. FREDDY
Wilderness Research Center, University of Idaho, Moscow, Idaho 83843
Present Address: Colorado Division of Wildlife, Box 481, Kremmling, Colorado 80459
Freddy, David J. 1979. Distribution and movements of Selkirk Caribou, 1972-1974. Canadian Field-Naturalist
93(1): 71-74.
Distribution and movements of Caribou inhabiting the Selkirk Mountains of northeastern Washington, northern Idaho, and
southern British Columbia were determined through aerial and ground surveys from 1972 to 1974. During most of the year,
Caribou were located in British Columbia at elevations above 1430 m, in close association with the spruce-fir forest.
Significant seasonal shifts in elevations frequented by these animals were not detected. Caribou consistently utilized specific
travel routes between and within drainages. Their only travel route to the USA was through Kootenay Pass. Continued
viability of the population probably depends on safeguarding their movement routes and spruce-fir forest winter ranges.
Key Words: Woodland Caribou, Selkirk Mountains, distribution, movements, ranges.
Woodland Caribou, Rangifer tarandus caribou
(Banfield 1961, p. 70), inhabiting the Selkirk Moun-
tains of northern Idaho, northeastern Washington,
and southern British Columbia (BC), Canada, may
constitute the only population of Caribou frequenting
the contiguous United States (USA). Caribou popula-
tions formerly existing in Maine and Minnesota were
extirpated during the early part of this century
(Palmer 1938; Nelson 1947). Populations in north-
western Montana and other portions of northern
Idaho have apparently been severely reduced in
number or extirpated (Evans 1960).
Historical reviews pertaining to Selkirk Caribou
were assembled by Evans (1960) and Layser (1974).
These accounts, based primarily on interviews, sum-
marize observations and records of these Caribou
from 1800 to 1971. Upon interpretation, these reviews
elucidate probable distribution and general move-
ments of this herd (Freddy 1974). Estimates of the
historic size of this population have been based on
inadequate data. Flinn (1959) felt there were at least
100 Caribou in the late 1950s while Layser (1974)
estimated that 15-20 animals existed in the early
1970s. Recent work indicates that 25-30 animals form
the population (D. Freddy. 1974. Report to inter-
national Caribou steering committee. Unpublished
report, B.C. Forest Service, Nelson. 9 pp.). Whether
this population has recently declined or is currently
increasing, decreasing, or stable in size is unknown.
In 1971 the University of Idaho Wilderness Re-
search Center and several organizations initiated a
cooperative study of Selkirk Caribou to document the
distribution and movements of this population. The
investigation was prompted by ongoing and potential
human activities that appeared to be threatening the
continued existence of these Caribou.
Study Area
The study was conducted in the Selkirk Mountains
between Priest Lake, Idaho and Kootenay Lake, BC.
(Figure 1). Elevations range from 700 m to 2100 m.
Maritime air from the Pacific Ocean continually
affects the area resulting in frequent cloudiness and
precipitation. Lower elevations receive about 75 cm
annually and higher elevations, over 250 cm of
precipitation, with 70% of this moisture occurring as
V2 THE CANADIAN FIELD-NATURALIST
snow between October and March (USDA 1972).
Snow begins to accumulate at higher elevations
during October and by December all elevations are
snow covered. In late February or March the
snowpack begins to settle and harden as warmer
temperatures become more frequent. During April,
snow at lower elevations begins to melt and at higher
elevations becomes thoroughly settled with depths of
3m common at elevations above 1500 m. By late
June, snow at lower elevations has melted but at
higher elevations snow will persist in sheltered areas
until early July.
Western Red-Cedar (Thuja plicata) — Western
Hemlock (T7suga heterophylla) and Engelmann
Spruce (Picea engelmanii) — Subalpine Fir (Abies
lasiocarpa) forest types dominate the area. The
spruce-fir forest predominates at elevations above
1500 m, spruce-fir and cedar-hemlock codominate at
1400-1500 m, while the cedar-hemlock forest domin-
ates at elevations below 1400 m. A more complete
description of these vegetative associations is pro-
vided by Daubenmire and Daubenmire (1968).
Primary land use has been and continues to be
timber production. British Columbia Highway 3
bisects the area and logging roads penetrate nearly
every drainage.
Methods
Distribution and movements of Caribou were
determined from aerial and ground observations of
Caribou or their tracks. Seventeen aerial surveys were
conducted from March 1972 through April 1973 using
a Cessna 182 while four surveys were conducted from
March to July 1974 with a Bell 206 helicopter. Flights
were conducted on days when visibility and contrast
were good and a snowpack was present. To avoid
duplicate observations of tracks, flights during winter
were made after recent snowfalls and during spring
after the snowpack had melted sufficiently to remove
tracks seen from previous flights. Fixed-winged
aircraft were flown at 135-140 km/h at altitudes
150-300 m above the ground, depending on terrain.
Helicopter surveys were generally flown at lower
altitudes and slower speeds. There was one observer
per flight. Aerial surveys followed fixed flight paths to
standardize data collection (Freddy 1974). These
Surveys concentrated on elevations above 1500 m
because Caribou and/or tracks could usually be seen
only on the snowpack within the relatively open-
crowned spruce-fir forest above this elevation. Mis-
identification of tracks during aerial surveys was
considered to be minimal. Moose (A/ces alces) and
Mountain Goat (Oreamnos americanus) were the
only other large ungulates present and the restricted
distributions of these two species minimized problems
in identifying tracks. Ground surveys were conducted
Vol. 93
from February 1972 through August 1974. These
surveys were used to supplement aerial surveys
especially in areas below 1500 m elevation. Areas
selected for search were judged probably to be utilized
by Caribou. Searches were made afoot and by
snowmobile.
Elevations where Caribou or tracks were located
were determined to the nearest 30 m from topo-
graphic maps and/or the altimeter of survey aircraft.
A one-way analysis of variance was used to test for
differences in monthly elevations of Caribou loca-
tions. Specific travel routes were determined by
tracking Caribou on foot or from aircraft.
Results and Discussion
From February 1972 through August 1974, 238
observations of Caribou or tracks were recorded.
Observations occurred during all months except
December and January of each year.
Caribou frequented areas within BC, Washington,
and Idaho. Areas of concentrated Caribou activity
occurred in BC with drainages immediately north of
Kootenay Pass receiving heavy utilization year round.
Areas in the USA most commonly frequented by
Caribou were adjacent to Snowy Top and Conti-
nental mountains (Figure |). Caribou were found in
BC and the USA during both winter and summer but
most of the population remained in BC year round.
This observed distribution of Caribou (Figure 1)
resembled the historic distribution of the population
(Freddy 1974). Current limits of distribution were
similar to historic bounds and areas adjacent to
Snowy Top and Continental mountains and Koot-
enay Pass continued as focal points of Caribou
activity. Areas in the USA within the southern portion
of the population’s range, however, apparently were
not used as intensely as in the past. This could have
been a function of the years of the study, rather than
an avoidance of areas in the USA. Skoog (1968, p.
437) and Kelsall (1968, p. 108) describe distributional
shifts of Caribou which can result in temporary
absence from areas.
Caribou were usually located year round at eleva-
tions above 1430 m with the mean elevation of Caribou
locations being 1657 + 227 (SD) m. There were no
significant differences in monthly elevation means
(P < 0.05) which ranged from 1591 + 204 (sD)
m in October to 1734 + 235 (Sp) m in September
(Table 1). The absence of distinct shifts in seasonal
elevations frequented by Selkirk Caribou differs from
movements reported for Caribou in Wells Gray
Provincial Park (WGPP) in central BC (Edwards and
Ritcey 1959). Edwards and Ritcey (1959) concluded
that the elevational distribution of WGPP Caribou
shifted markedly in response to accumulating or
receding snowpacks. During fall, Caribou in WGPP
1979
he @
MTN.
COMMON
4
1
t]
DISTRIBUTION \
i]
\
PRIEST
OCCASIONAL LAKE
INFREQUENT
MOVEMENT
ROUTE
HIGHWAY
FIGURE 1. Distribution, frequency of occurrence, and
known movement routes of Selkirk Caribou, 1972-1974.
moved down into the cedar-hemlock forest to avoid
deepening snow at higher elevations and to forage on
understory vegetation. As snow accumulated, Cari-
bou ascended to the spruce-fir forest to forage on
arboreal lichens, and then again descended to lower
elevations in the spring. Although movements of
Selkirk Caribou were apparently more variable, both
populations frequented lower elevations in fall and
associated with the spruce-fir forest in mid-winter.
Food habits of Selkirk and WGPP Caribou (Freddy
TABLE |—Elevations at which Caribou or their tracks were
observed, 1972-1974
Number of Elevation (m),
Month * observations Mean +SD Range
February 22 1601 + 234 976-1891
March 24 1712 + 229 976-1952
April 19 1674 + 169 1281-1952
May 56 1677 + 185 946-1952
June 27 1665 + 154 1342-1891
July D2 1693 + 210 1312-2074
August 21 1638 + 176 1281-2044
September 14 1734 + 235 1281-2013
October 20 1591 + 204 1281-1952
November 13 1645 + 181 1342-1952
Totals 238 OS ae DLT) 946-2074
a .
Surveys were not conducted in December or January.
NOTES 73
1974: Edwards and Ritcey 1960) also suggest similar
responses to changing snow conditions and availa-
bility of forage.
Caribou consistently followed specific travel routes
between and within drainages (Figure 1). Routes
commonly incorporated natural passes along ridges,
frequently followed stream bottoms, invariably pro-
ceeded through forested areas, and generally con-
nected feeding and resting areas used by Caribou.
Most routes were utilized during all seasons.
Critical to international movement of animals was
the travel route through Kootenay Pass where
Caribou had to cross a major highway (Figure 1).
Kootenay Pass was the only known route used by
Caribou to enter the USA, and historical accounts
(Spry 1968, p. 480; Layser 1974) support the impor-
tance and long-term use of this route by Caribou.
Elevations frequented by Selkirk Caribou indicate
that these animals utilize the cedar-hemlock forest
and especially the spruce-fir forest year round.
Forested areas above 1430 m must be considered
vitally important to this population. The close
association of Selkirk Caribou to the spruce-fir forest
during winter reflects their use of arboreal lichens
(Alectoria spp.) as a primary winter food (Freddy
1974). Edwards and Ritcey (1960) also found arboreal
lichens to be a principal winter food for Caribou in
WGPP. Edwards et al. (1960) indicated that mature
stands of spruce-fir, as opposed to cedar-hemlock,
produced the greater amount of arboreal-lichen
biomass available for consumption by Caribou. The
association of A/ectoria spp. with mature stands of
spruce-fir places this forage in conflict with extensive
clearcut logging (Edwards et al. 1960; T. Ahti. 1962.
Ecological investigations on lichens in Wells Gray
Provincial Park, with special reference to their
importance to mountain caribou. Unpublished re-
port, Department of Botany, University of Helsinki,
Finland. 63 pp.).
Consistent use by Caribou of specific travel routes
implies that routes should remain undisturbed to
allow established patterns of distribution and range
use to continue. Klein (1971) reported that Reindeer
(R. t. tarandus) have strong traditions for specific
migratory routes and that realignment of such routes
is difficult. Proposed expansion of BC Highway 3 to
four lanes at Kootenay Pass could easily jeopardize
future movements of Caribou into the USA. Klein
(1971) reported highways and railroads have ob-
structed movements of wild reindeer in Norway.
The continued viability of the Selkirk Caribou
population depends on cooperative international
management. Although Caribou continue to frequent
the northwestern USA, in all likelihood perpetuation
of mature spruce-fir forests frequented by Caribou in
BC is essential to the occurrence of these animals in
74 THE CANADIAN FIELD-NATURALIST
the USA. Timber harvest programs and other human
developments in both BC and the USA must be
coordinated to maintain known movement routes and
spruce-fir winter ranges.
Acknowledgments
Financial support for the work was received from
the British Columbia Fish and Wildlife Branch,
British Columbia Forest Service, Idaho Department
of Fish and Game, Idaho Wildlife Federation—
District 1, Inland Empire Big Game Council, National
Rifle Association, National Wildlife Federation,
Priest Lake Sportsmen’s Association, Sierra Club
Foundation, US Forest Service—Region |, Washing-
ton Department of Game, Washington State Sports-
men’s Council, West Kootenay Outdoorsmen, and the
University of Idaho. A. W. Erickson and D.R.
Johnson are gratefully acknowledged for initiating
the study and obtaining financial support. G. Koehler
was invaluable as a field assistant during winter. L. H.
Carpenter, R. B. Gill, D.R. Johnson, and D.R.
Miller read the manuscript.
Literature Cited
Banfield, A. F. 1961. A revision of the Reindeer and
Caribou, genus Rangifer. National Museum of Canada
Bulletin 177. 137 pp.
Daubenmire, R. and J. Daubenmire. 1968. Forest vegeta-
tion of eastern Washington and northern Idaho. Washing-
ton Agricultural Experiment Station Technical Bulletin
60. 104 pp.
Edwards, R. Y. and R.W. Ritcey. 1959. Migrations of
Caribou in a mountainous area in Wells Gray Park,
British Columbia. Canadian Field-Naturalist 73(1):
21-25.
Edwards, R. Y. and R. W. Ritcey. 1960. Foods of Caribou
Vol. 93
in Wells Gray Park, British Columbia. Canadian Field-
Naturalist 74(1): 3-7.
Edwards, R.Y., J. Soos, and R.W. Ritcey. 1960.
Quantitative observations on epidendric lichens used as
food by Caribou. Ecology 41(3): 425-431.
Evans, H. F. 1960. A preliminary investigation of Caribou
in northwestern United States. M.Sc. thesis, Montana
State University, Bozeman. 145 pp.
Flinn, P. 1959. The Caribou of northern Idaho. Idaho
Wildlife Review 11(5): 10-11.
Freddy, D. J. 1974. Status and management of the Selkirk
Caribou herd, 1973. M.Sc. thesis, University of Idaho,
Moscow. 132 pp.
Kelsall, J. P. 1968. The migratory Barren-ground Caribou
of Canada. Canadian Wildlife Service Monograph
Number 3. 340 pp.
Klein, D. R. 1971. Reactions of Reindeer to obstructions
and disturbances. Science 173: 393-398.
Layser, E. F. 1974. A review of the mountain Caribou of
northeastern Washington and adjacent northern Idaho.
Journal of the Idaho Academy of Science, Special
Research Issue 3. 63 pp.
Nelson, E. C. 1947. The Woodland Caribou in Minnesota.
Journal of Wildlife Management 11(3): 283-284.
Palmer, R.S. 1938. Late records of Caribou in Maine.
Journal of Mammalogy 19(1): 37-43.
Skoog, R.O. 1968. Ecology of the Caribou (Rangifer
tarandus granti) in Alaska. Ph.D. thesis, University of
California, Berkeley. 699 pp.
Spry, I. M. (Editor). 1968. The papers of the Palliser
expedition 1857-1860. Champlain Society, Toronto. 694
pp.
U.S. Department of Agriculture. 1972. Kaniksu National
Forest multiple use plan. Kaniksu National Forest,
Sandpoint, Idaho. 50 pp.
Received 9 January 1978
Accepted 10 August 1978
Effects of Fire on the Location of a Sharp-tailed Grouse Arena
DONALD A. SEXTON! and MURRAY M. GILLESPIE2
'Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2
*Department of Renewable Resources and Transportation Services, Box 14, 1495 St. James Street, Winnipeg, Manitoba
R3H 0W9
Sexton, Donald A. and Murray M. Gillespie. 1979. Effects of fire on the location of a Sharp-tailed Grouse arena.
Canadian Field-Naturalist 93(1): 74-76.
Fire is known to be an important factor in creating
and maintaining Sharp-tailed Grouse (Pedioecetes
Phasianellus) habitat (Amman 1957; Kirsch and
Kruse 1972). Controlled burning has been used to
improve Sharp-tailed Grouse habitat in several areas
(Amman 1957, 1963; Baumgartner 1939; Grange
1948: Kirsch and Kruse 1972; Miller 1963). Few
observations are available, however, on the im-
mediate effects of fire on Sharp-tailed Grouse
behavior. Amman (1957) suggests that fire does not
affect arena use by males. Anderson (1969) reports
male Greater Prairie Chicken ( 7ympanuchus cupido)
1979
continuing to use an arena the day after it had been
burned. During the spring of 1975 we observed the
effects of a fire on a Sharp-tailed Grouse arena near
Chatfield, Manitoba (50°47’'N, 97°34’W).
Since 1970, spring surveys of arenas have been
conducted in the Chatfield area. Known arenas were
visited in early morning and numbers of male and
female grouse recorded. The counts were conducted
three or more times a week from early April until mid-
June.
Arena A was used by males in 1971 but not in 1972
or 1973. Surveys were not conducted in 1974 owing to
flooding of much of the area. During the spring of
1975, a new arena (B) was located 480 m north of the
now abandoned arena A, in the same area of open
grassland (Figure |). Vegetation in the grasslands (Gr)
consisted of native grasses, chiefly Needle Grass
(Stipa spartea), June Grass (Koeleria cristata), and
wheat grasses (Agropyron spp.) as well as a variety of
forbs and scattered Saskatoon (Amelanchier alni-
folia) and Snowberry (Symphoricarpos occidentalis)
shrubs. The shrub area (Sh) contained clumps of
Dwarf Birch ( Betula glandulosa) and Saskatoon. The
forested portion (Fo) was dominated by Trembling
Aspen (Populus tremuloides).
When arena B was discovered on7 May, 10 birds of
unknown sex were present. Counts on subsequent
mornings showed between 7 and 10 males displayed at
this arena prior to 23 May. During this period birds
were never observed displaying at arena A. On 23 and
24 May, a fire burned the southwestern portion of the
grassland area and eliminated all residual grass and
forbs but did not greatly affect shrub or tree cover in
the area burned. Arena A was completely burned over
whereas arena B was unaffected (Figure 1).
On 25 May two non-displaying Sharp-tailed
Grouse were present on arena A and eight males were
displaying on arena B. Between 26 May and | June,
one or two birds were displaying on arena A, and four
to seven on arena B each morning. By 16 June only
one bird remained on arena B while arena A had five
to eight males actively displaying. Other neighboring
arenas showed a decrease in the number of males
during June, as is typical in late spring (Hamerstrom
1939). On 17 June no birds were on arena B but five
birds were still attending arena A. After this date birds
were not present on either arena.
The difference in vegetative cover between arenas A
and B prior to the fire was not obvious. After the fire
arena B was still covered by rank residual grass and
forbs in addition to the current season’s growth.
Arena A had only new growth which was sparse until
several weeks after the fire.
Several authors have shown that most Sharp-tailed
Grouse arenas are in areas of sparse ground cover
(Amman 1957; Kobridger 1965; Sisson 1975). Ac-
NOTES 15
FIGURE 1.
(++) on Chatfield study area. Note grassland (Gr),
shrub (Sh), and forest (Fo).
Location of arenas and extent of burned area
cording to Amman (1957), males may abandon sites
with tall or dense ground cover. The shift we observed
at Chatfield may be explained by birds moving back
to the traditional arena (A) after its vegetation cover
was reduced by fire to a more acceptable height. The
lack of tall cover would permit displaying males to see
and be seen, which are two requirements of arena sites
(Anderson 1969). In 1976 and 1977, displaying males
used arena A regularly but were never seen where
arena B had been in 1975.
These observations were made during a study of
population, movements, and habitat use of Sharp-
tailed Grouse conducted by the Manitoba Depart-
ment of Renewable Resources and Transportation
Services. We thank G. W. Pepper, S. G. Sealy, and
M. W. Shoesmith for comments on the manuscript.
76 THE CANADIAN FIELD-NATURALIST
Literature Cited
Amman, G. A. 1957. The prairie grouse of Michigan.
Michigan Department of Conservation Technical Bul-
letin, Lansing. 200 pp.
Amman, G.A. 1963. Status and management of Sharp-
tailed Grouse in Michigan. Journal of Wildlife Manage-
ment 27: 802-809.
Anderson, R. K. 1969. Prairie chicken responses to chang-
ing booming-ground cover type and height. Journal of
Wildlife Management 33: 636-643.
Baumgartner, F.M. 1939. Studies in distribution and
habits of the Sharp-tailed Grouse in Michigan. Trans-
actions of the North American Wildlife Conference 4:
485-490.
Grange, W. B. 1948. Wisconsin grouse problems. Wiscon-
sin Conservation Department Publication 328. 318 pp.
Vol. 93
Hamerstrom, F. N., Jr. 1939. A study of Wisconsin Prairie
Chicken and Sharp-tailed Grouse. Wilson Bulletin 51:
105-120.
Kirsch, L. M.and A. D. Kruse. 1972. Prairie fires and wild-
life. Proceedings of the Tall Timbers Fire Ecology Con-
ference 12: 289-303.
Kobridger, G. D. 1965. Status, movements, habitats and
food of Prairie Grouse on a Sandhills Refuge. Journal
of Wildlife Management 29: 788-800.
Miller, H. A. 1963. Use of fire in wildlife management.
Proceedings of the Tall Timbers Fire Ecology Conference
2: 127-143.
Sisson, L. 1975. The Sharp-tailed Grouse in Nebraska.
Nebraska Game and Parks Commission. 88 pp.
Received 13 February 1978
Accepted 6 August 1978
Brewer’s Blackbird Breeding in the Northwest Territories
PHILIP H. R. STEPNEY
Ornithology Department, Provincial Museum of Alberta, Edmonton, Alberta T5N 0M6
Stepney, Philip H.R. 1979. Brewer’s Blackbird breeding in the Northwest Territories. Canadian Field-Naturalist
93(1): 76-77.
The occurrence of Brewer’s Blackbird breeding in the Northwest Territories was verified at Fort Simpson, 27 June 1977.
Additional courting individuals were observed at Hay River on two occasions in the same year. This breeding record
represents an appreciable northward expansion of the range of this species and is seemingly related to increased modification
of previously forested areas.
Key Words: Brewer’s Blackbird, breeding, range extension, Northwest Territories.
Brewer’s Blackbird (Euphagus cyanocephalus) has
increased both its breeding range (Stepney and Power
1973) and winter range (Stepney 1975) inthis century.
Typically distributed in western North America,
(A.O.U. 1957), habitat modification resulting from
agricultural and transportation route development
has seemingly aided the expansion of this species
(Roberts 1932).
Previously, the Meikle River formed the northern
limit of the breeding range of Brewer’s Blackbird in
the Peace River region of Alberta (Godfrey 1966). In
adjacent northeastern British Columbia, a “family”
group had been recorded in 1967 at Fort Nelson,
where the birds were observed again in 1974 (Erskine
and Davidson 1976). Previous authors had not
recorded this species in the Fort Nelson region. There
are apparently three earlier records of non-breeding
Brewer’s Blackbirds occurring in the territories: Fort
Simpson, District of Mackenzie, on 15 June 1958,
Baker Lake, District of Keewatin, in November 1923
(Godfrey 1966), and north of Old Crow, Yukon
Territory, (three birds) in 1971 (Schweinsburg 1974).
In 1977 the author observed Brewer’s Blackbirds in
two locations in the District of Mackenzie. At Hay
River, on 28 May, two pairs of mated birds were
observed along the railroad right-of-way adjacent to
the airfield. The males were observed courting and
guarding the females and engaging in pursuit flights,
behavior typical of breeding Brewers Blackbirds
(Williams 1952). Time did not permit a search for
nests. On 8 September 1977 Brewer’s Blackbirds were
again recorded in Hay River. Four mature males were
observed within the town, suggesting that a small
colony was present in Hay River that summer. At Fort
Simpson, on 27 June, a single pair of adults with five
fledglings were observed in the cleared area adjacent
to the gravel airstrip within the town. The young were
flying among the willow shrubs paralleling the
airstrip; the adults mobbed me while I searched
unsuccessfully for the nest-site, which probably was
located on the grass-covered bank of the airstrip
drainage ditch.
The occurrence of Brewer’s Blackbirds at Fort
Simpson is approximately 360 km northeast of their
1979
previous northern limit in British Columbia. This
breeding expansion is appreciable but not unexpected
in view of the highway and rail-line developments in
this originally forested area. It seems likely that the
modified habitats along these rights-of-way provided
dispersal routes for Brewer’s Blackbird. This species
has been observed penetrating forested areas along
cleared rights-of-way in areas of Alberta (Smith
1975), Saskatchewan and British Columbia (A. J.
Erskine, personal communication).
I acknowledge the assistance of Renewable Re-
sources Consulting Services Ltd., with whom I was
employed at the time of these observations.
Literature Cited
American Ornithologists’ Union. 1957. Check-list of North
American birds. Fifth edition. American Ornithologists’
Union.
Erskine, Anthony J. and Gary S. Davidson. 1976. Birds in
the Fort Nelson lowlands of northeastern British Colum-
bia. Syesis 9: 1-11.
NOTES 77
Godfrey, W.E. 1966. The birds of Canada. National
Museum of Canada, Bulletin Number 203, Biological
Series Number 73. 428 pp.
Roberts, T. S. 1932. The birds of Minnesota: Volume 2.
University of Minnesota Press, Minneapolis.
Schweinsburg, R. E. 1974. An ornithological study of pro-
posed gas pipeline routes in Alaska, Yukon Territory and
the Northwest Territories, 1971. Arctic Gas Biological
Report Series 10: 1-157.
Smith, H. C. 1975. The birds north of Lesser Slave Lake,
Alberta. Blue Jay 33: 232-239.
Stepney, Philip H.R. 1975. Wintering distribution of
Brewer’s Blackbird: historical aspect, recent changes, and
fluctuations. Bird-Banding 46: 106-125.
Stepney, P. H.R. and Dennis M. Power. 1973. Analysis
of the eastward breeding expansion of Brewer’s Black-
bird plus general aspects of avian expansions. Wilson
Bulletin 85: 452-464.
Williams, L. 1952. Breeding behavior of the Brewer Black-
bird. Condor 54: 3-47.
Received 6 April 1978
Accepted 21 September 1978
A Trap to Measure Populus and Salix Seedfall
JOHN C. ZASADA! and ROSEANN DENSMORE2
'USDA Forest Service, Institute of Northern Forestry, Fairbanks, Alaska 99801
2Botany Department, Duke University, Durham, North Carolina 27706
John C. Zasada and Roseann Densmore. 1979. A trap to measure Populus and Salix seedfall. Canadian Field-
Naturalist 93(1): 77-79.
A seed trap, a wooden frame(1 X 1 m X 5 cm) witha plastic sheeting on the bottom to hold moistened potting soil was used in
Alaska to determine seedfall for Populus spp. (poplars) and Salix spp. (willows) through germination of seedlings. This seed
trap takes advantage of these species’ ability to germinate rapidly under ambient temperatures and optimum moisture
conditions.
Key Words: Seed dispersal, natural regeneration, seed trap, Salicaceae, Alaska.
The types of seed traps used to measure seedfall
vary greatly. Forest tree seedfall is usually measured
with rectangular or circular traps which vary in
surface area and height of collecting surface above
ground (Sarvas 1962; Zasada and Gregory 1972).
Werner (1975) described a trap that contained a sticky
surface to which seeds adhered. Ryvarden (1971) used
a water-filled trap to study seed dispersal in alpine
areas of Norway.
Although these traps could be used for studying
Populus (poplar) and Salix (willow) seedfall in
Alaska, two factors make them less than perfect. First,
within any part of Alaska, there may be more than one
species from each genus contributing to the seedfall.
Although these species disperse seed at different
times, there can be significant overlap. Thus it is
desirable to identify the contribution of each species.
Seeds of the two genera can be separated on the basis
of color (i.e., Populus seed is tan to white in color,
Salix seed is green). Further, it is possible after some
experience to separate P. tremuloides (Trembling
Aspen) from P. balsamifera (Balsam Poplar). Willow
seeds, however, cannot be separated easily on the
basis of features identifiable in the field. Second,
because of the small size of these seeds it is laborious
and time-consuming to separate seeds from other
debris that lands in seed traps and then to germinate
them.
We designed, and have been using for 2 yr, a seed
trap that takes advantage of the rapid germination of
78 THE CANADIAN FIELD-NATURALIST
Vol. 93
FIGURE |. Seed trap with aspen seedlings in early July. Toothpicks mark seedling locations.
Populus and summer-dispersing Salix under opti-
mum conditions of temperature and moisture (Zasada
and Viereck 1975). Seeds.of these two genera must
germinate shortly after landing on the seedbed or they
die (Schreiner 1974; Brinkman 1974).
Our trap is a I- X I-m wooden frame (5 cm deep)
covered on the underside with one layer of 4-mil
plastic sheeting (Figure 1). The trap 1s filled with a
standard potting soil, such as Jiffy-Mix,! that is kept
wet. If flooding from heavy rains is a problem, holes
should be cut in the frame just below soil level. So that
soil stays wet, holes should not be punched in the
sheeting.
It is important that the commercial potting mixes
not have an inhibitory effect on germination of these
species. We have not observed any effect from Jiffy-
Mix! or Jiffy-Mix Plus! in growth chamber studies.
Traps should be placed in the field prior to seed
dispersal. Germinants are counted periodically and
left to develop until they can be identified. Seed
collected from identified willows and poplars and
germinated on trap soils can be used as an aid to
identify unknown germinants.
We have used the traps on upland and floodplain
sites in interior Alaska. Ona birch (4.9 ha) and aspen
(2.8 ha) clearcut we used four traps in each area to
measure seedfall. Aspen seedfall was estimated to be
383 seeds/ m2 in the aspen clearcut and 376 seeds/ m2
in the birch clearcuts (coefficients of variation 25%
'The use of trade, firm, or corporation names does not
constitute endorsement by the U.S. Department of Agri-
culture.
and 19%, respectively). The first seed dispersal was
observed on 10 June, the first germinants between 12
and 16 June, and the last germinants on 14 July (Table
1). Only aspen seedlings were observed in these traps.
Figure | shows traps in early July.
Four traps placed on a floodplain produced
seedlings of a number of species (Table 2). The traps
were set out after dispersal of Salix alaxensis. They
were within an 8-yr-old stand (traps | and 2) or onan
open sandbar adjacent to it (traps 3 and 4). The main
species in this stand were S. interior, S. alaxensis, and
P. balsamifera;, less common species were S. novae-
anglicae and S. monticola. Seedlings of species that
are rare or absent on these sites (e.g., S. bebbiana, P.
tremuloides) were produced as well as those of species
occurring in the stand.
TABLE |—Number of aspen germinants and percent of total
number germinating on a given date in four seed traps
located in clearcuts on upland sites in 1977
Aspen Birch
Number of
germinants
Mean + sD 383 +99 370 22 7/2)
(range) (277-S 16) (353-436)
Percent
germinating
16 June 3p).3) 46.8
23 June 26.2 36.7
3 July 7.8 dil
7 July 9.6 7.5
14 July let Hes)
79
NOTES 79
TABLE 2—Species composition and number of germinants observed in four seedtraps located on floodplain sites adjacent
to the Tanana River, Alaska
Traps within 8-yr-old stand Traps on open sandbar
Species | 2 3 4
Salix lasiandra' 12 5 2
S. bebbiana 33 50 5 12
S. arbusculoides 3 2 = =
S. monticola 5 0 = —
S. interior 45 281 7 26
S. novae-anglicae 6 2 | I
S. planifolia I 2B l |
Populus balsamifera 4 2 5 5
P. tremuloides 41 49 48 3
Unknown Salix sp. 14 4 9 3
'Terminology follows Argus (1973).
Our observations demonstrate that the seed trap
is feasible, but several drawbacks exist. First, depen-
ding on the weather conditions, the traps may have to
be watered as frequently as every day. Second, the
variable measured with these traps is the number of
germinants, which is not necessarily the total number
of potentially viable seeds reaching the trap. We
believe that by keeping the traps saturated, the
number of germinants is a good estimate of the
number of viable seeds; but this point needs further
examination.
Literature Cited
Argus, George W. 1973. The genus Sa/ix in Alaska and the
Yukon. National Museum of Natural Sciences, National
Museum of Canada, Publications in Botany Number 2.
279 pp.
Brinkman, Kenneth A. 1974. Salix L. In Seeds of woody
plants in the United States. USDA Agriculture Hand-
book Number 450. pp. 746-750.
Ryvarden, L. 1971. Studies in seed dispersal. I. Trapping of
diaspores in the alpine zone of the Finsé, Norway.
Norwegian Journal of Botany 18: 215-226.
Sarvas, Risto. 1962. Investigations on the flowering and
seed crop of Pinus silvestris. Finnish Forest Research
Institute Publication 53.4. 198 pp.
Schreiner, Ernst J. 1974. Populus L. In Seeds of woody
plants in the United States. USDA Agriculture Handbook
Number 450. pp. 645-655.
Werner, P. A. 1975. A seed trap for determining patterns of
seed deposition in terrestrial plants. Canadian Journal of
Botany 53: 810-813.
Zasada, John C. and Robert A. Gregory. 1972. Paper birch
seed production in the Tanana Valley, Alaska. USDA
Forestry Service Research Note PNW-177. Pacific North-
west Forest and Range Experiment Station, Portland
Oregon. 7 pp.
Zasada, John C. and Leslie A. Viereck. 1975. Effect of
temperature and stratification on germination of Alaskan
Salicaceae. Canadian Journal of Forestry Research 5:
333-337.
Received 7 April 1978
Accepted 19 August 1978
Examination of Overwintering Adult Carabid Beetles
for Associated Mites
J. E. OLYNYK and R. FREITAG
Department of Biology, Lakehead University, Thunder Bay, Ontario P7B S5El
Olynyk, J. E. and R. Freitag. 1979. Examination of overwintering carabid beetles for associated mites. Canadian Field-
Naturalist 93(1): 79-81.
Mites are commonly associated with adult carabids in northwestern Ontario from spring to fall but are not normally found
on the beetles during winter. This suggests that the mites are phoretic on the carabids, using them for transport during times of
the year when the carabids are active.
Key Words: northwestern Ontario, mites, beetles, Carabidae, Acari, ectoparasites.
80 THE CANADIAN FIELD-NATURALIST
In North America, collections of carabids during
winter have been rare and no previous work has been
done on mite populations associated with hibernating
carabids. Aitchison (1976), using pitfall traps, studied
numerous invertebrate groups in subnivean environ-
ments in southern Manitoba. She collected very few
carabids in December and March and none in
January or February. Olynyk and Freitag (1977) did
not capture any carabids in pitfall traps near Thunder
Bay, Ontario, between 9 November and 6 April,
though Olynyk (1978) found an average of approxi-
mately eight mites per captured carabid between May
and October. Larochelle (1972a, b, 1973, 1974) made
several collections of hibernating adult carabids in
southern Quebec but did not record the presence of
mites on the beetles (personal communication).
The purpose of this study was to determine winter
associations of ground beetles and mites in the context
of annual relationships.
Materials and Methods
Carabids were collected between 6 and 26 April
1977 in 200 pitfall traps near Thunder Bay, Ontario
(Olynyk and Freitag 1977). Three hand collections
were made in the same location in 1977 on 19
November and 3 and 23 December. Mounds of earth,
rotten stumps and logs, and surrounding areas on, in,
and under the leaf litter were inspected. Larochelle
(1974, 1975) noted that such locations were some of
the most productive sites for collecting carabids in
winter. On 22 April 1978, hand collections were made
beside Ontario Highway 588 near the junction of
Highways 11/17 and 588 in northwestern Ontario.
Areas under loose materials including logs, metal,
tires, boards, and cardboard were inspected.
All carabids collected were placed in individual
vials of 70% ethanol. External surfaces and areas
under elytra were examined on each beetle collected
under a Wild MS dissecting microscope. Alcohol from
vials containing carabids was filtered through Kim-
wipe tissue and the tissue then examined for mites.
Carabids were identified to species using identifica-
tion keys by Lindroth (1961-1969).
Results and Discussion
Table | contains a list of the carabids collected. On
the 53 carabids collected, representing five genera and
Six species, we found no mites on external surfaces or |
under elytra.
Carabids collected in November and December
were found beneath rotten stumps and were torpid.
Approximately 80% of the carabids collected on 22
April were found beneath a sheet of metal, the rest
beneath a nearby sheet of cardboard. Burrows were
present, the carabids were congregated and many
were torpid. The ground was frozen.
Vol. 93
TABLE 1—Carabids collected in early spring and winter of
1977 and 1978 in pitfall traps and by hand near Thunder
Bay, Ontario
Sex
Date Carabid species on 9
26 April 1977 Prerostichus
adstrictus l 3
P. pensylvanicus 4 7
19 November 1977 Sphaeroderus
nitidicollis I 2
Calosoma frigidum l
P. pensylvanicus 2
Agonum decentis ]
3 December 1977 S. nitidicollis l
23 December 1977 S. nitidicollis I
22 April 1978 P. pensylvanicus | 3
Calathus ingratus 1] 14
Greene (1975) found acarid hypopi appearing on
carabids in August in Washington, with many
hundreds commonly present on the carabids by
October. No data were recorded for hibernating
carabids, but few if any mites were present on carabids
in spring, which suggests that the mites did not
overwinter on the carabids. W.M. Graham of
Lakehead University studied ants and associated
mites populations near Thunder Bay in 1969 and 1970
(March 1978, personal communication). Average
mite numbers per ant were lowest in spring and fall,
suggesting that the mites did not overwinter on the
hibernating ants. Olynyk (1978) noted that the
numbers and frequency of occurrence of mites on
carabids were lower in spring and fall. As none of the
53 hibernating adult carabids examined carried mites,
it can be assumed that the mites do not overwinter on
the carabids.
Aitchison (1976) found that carabids in southern
Manitoba were inactive at temperatures below about
-2°C. She collected large numbers of mites in mid-
winter and concluded that their activity seemed to be
unrestricted by temperatures encountered in her
study.
Mites phoretic on carabids, using them solely as a
transport mechanism, would gain no such benefit
from hibernating carabids. It follows that mites
phoretic on carabids in other regions likewise are not
found on hibernating carabids.
Acknowledgments
We are indebted to J. P. Walas for his aid in
collecting specimens and to the National Research
Council of Canada for financial support (Grant No.
A4888).
IDG)
Literature Cited
Aitchison, C. W. 1979. The activity of subnivean inverte-
brates in Southern Manitoba. M.Sc. thesis, University of
Manitoba, Winnipeg, Manitoba. 197 pp.
Greene, A. 1975. Biology of five species of Cychrini
(Coleoptera: Carabidae) in the steppe region of South-
eastern Washington. Melanderia 19: 1-43.
Larochelle, A. 1972a. Collecting hibernating ground
beetles in stumps. (Coleoptera: Carabidae). Coleopterists’
Bulletin 26(1): 30.
Larochelle, A. 1972b. Collecting hibernating Carabidae
under snow. Proceedings of the Entomological Society of
Washington 74(4): 473.
Larochelle, A. 1973. Collecting hibernating ground beetles
under snow. Entomological News 84: 82.
Larochelle, A. 1974. Winter habits of carabid beetles
NOTES 81
(Coleoptera: Carabidae). Great Lakes Entomologist 7(4):
143-145.
Larochelle, A. 1975. La chasse aux Carabidae en hiver.
Cordulia 1(4): 117-118.
Lindroth, C. H. 1961-1969. The ground-beetles of Canada
and Alaska. Opuscula Entomologica Supplementum.
1192 pp.
Olynyk, J. E. 1978. Carabidae (Coleoptera) and associated
Acari in Northwestern Ontario. M.Sc. thesis, Lakehead
University, Thunder Bay, Ontario. 73 pp.
Olynyk, J. and R. Freitag. 1977. Collections of spiders
beneath snow. Canadian Field-Naturalist 91(4): 401-402.
Received 15 May 1978
Accepted 29 August 1978
Ring Counts in Salix arctica from Northern Ellesmere Island
D. B. O. SAVILE
Biosystematics Research Institute, Central Experimental Farm, Ottawa, Ontario KIA 0C6
Savile, D. B. O. 1979. Ring counts in Salix arctica from northern Ellesmere Island. Canadian Field-Naturalist 93(1):
81-82.
In 1962 I collected two large specimens of Salix
arctica Pall., which had been eroded out of steep-
sided postglacial benches roughly 100 m above the
present level of Lake Hazen, Northwest Territories, at
about 81°50’N, 71°15’W. They were to have been
picked up by R. E. Beschel, but, when he failed to do
so, they lay forgotten in a drawer for years.
I have now cut sections from the main stem and a
major branch of each specimen. The sections were
polished, cleared with lacquer, and ring counts were
made under the dissecting microscope. The short cool
growing season in the high Arctic makes some rings
very narrow or incomplete; and some such rings may
not be definitely distinguishable even under the
microscope.
The larger plant (Savile 4841A) yielded counts of
84, 84, and 85 (at least partial) rings on a radius of
about 36 mm from the markedly eccentric origin.
These are minimum counts, because some of the early
rings are thought to have included some with almost
zero growth. The outermost rings are relatively wide,
indicating vigorous growth in the final years of the life
of the plant, A large branch yielded 36 rings without
any wide outer ones. Possibly death of this branch
diverted nutrients into the main stem. R. E. Beschel
and D. Webb (1964. Axel Heiberg Island Preliminary
Report. Edited by F. Muller. McGill University,
Montreal. pp. 189-198. Processed) indicate that S.
arctica periodically loses branches.
The second specimen (Savile 484/ B), with maxi-
mum wood radius of 28 mm, yielded a minimum of
43,44, 44, and doubtfully 48 rings in separate sections,
and again most of the last few rings were relatively
wide. Sections from both specimens have been
deposited in DAO.
I suspect that both plants were killed by erosion of
the bench on which they grew, because vigorous
growth in their last years does not suggest the
senescence inferred by Warren Wilson (1964) for
plants of this species at Resolute (74°41’N), where
little or no growth seemed to occur after about SO yr.
Although the Hazen Camp specimens are from about
81°50’N, they are better grown than any at Resolute,
largely because Hazen Valley, being ringed by
mountains, is subject to dynamically warmed winds
and has relatively high summer temperatures; but
partly, perhaps, because sites such as the raised
branches have good drainage and a deep active layer.
The difference is reflected in the flora: about 115
species of vascular plants in Hazen Valley and 70 at
82 THE CANADIAN FIELD-NATURALIST |
Resolute. For comparison of these and some other
arctic sites see Savile (1972, p. 11). It should be noted
that on some sites, notably depressions flooded by
Lake Hazen in midsummer, Salix arctica makes very
poor growth.
It seems probable that, barring soil movement or
other accidents, plants on favorable sites in Hazen
Valley have either an indefinite life or a much higher
life limit than at Resolute. It is worth noting that, of
the 12 plants counted from Axel Heiberg Island (all
about 79°25’ to 79°30’N) by Beschel and Webb, one
showed 87 and all the others between 18 and 64 rings.
My specimen, 484/ A, is perhaps the oldest recorded
north of 80°N. A few large and vigorous plants up to
about 3-m spread were seen near Hazen Camp, but I
did not feel justified in sacrificing any of them for the
Vol. 93
sake of a ring count.
I am grateful to a reviewer of this manuscript for
drawing my attention to the mention by Raup (1965)
of specimens of Salix arctica at about 73°N in
Northeast Greenland of up to 210 and 236 yr old.
Literature Cited
Raup, H. M. 1965. The structure and development of turf
hummocks in the Mesters Vig district, Northeast Green-
land. Meddelelser om Gronland 165(3): 1-112.
Savile, D. B. O. 1972. Arctic adaptations in plants. Canada
Department of Agriculture, Research Monograph 6.
Warren Wilson, J. 1964. Annual growth of Salix arctica
in the high Arctic. Annals of Botany (London) 28: 71-76.
Received 4 July 1978
Accepted 5 September 1978
Giant Cow Parsnip (Heracleum mantegazzianum) on
Vancouver Island, British Columbia
NEIL K. DAWE! and ERIc R. WHITE?
'Canadian Wildlife Service, R.R. 1, Qualicum Beach, British Columbia VOR 2T0
°5312 Ewart Street, Burnaby, British Columbia V5J 2W4
Dawe, Neil K. and Eric R. White. 1979. Giant Cow Parsnip ( Heracleum mantegazzianum) on Vancouver Island, British
Columbia. Canadian Field-Naturalist 93(1): 82-83.
The known distribution of the Giant Cow Parsnip ( Heracleum mantegazzianum) on Vancouver Island, British Columbia is
reported. A discussion of the time span the plant has taken to become established is included along with notes on the dispersal
of the plant from one site of introduction.
Key Words: distribution, Heracleum mantegazzianum, Vancouver Island, weeds.
The distribution of the Giant Cow Parsnip (Her-
acleum mantegazzianum) in Canada was recently
described by Morton (1978) as occurring primarily in
southern Ontario. He also cites a record from
Vancouver on the west coast (Kamermans 1977):
however, there is no mention of the plant’s occur-
rence on Vancouver Island. Taylor and MacBryde
(1977) list but one species of Heracleum for British
Columbia: H. sphondylium (= H. lanatum). This note
documents the occurrence of H. mantegazzianum on
Vancouver Island and will alert naturalists to the
possible occurrence of the species in other areas of the
province.
During summer field work of 1978 we discovered a
large number of H. mantegazzianum plants growing
along the banks of French Creek, near the creek
mouth, approximately 6 km NW of Parksville, British
Columbia (49°21’N, 124°22’W). Subsequently two
other groups of plants of that species were brought to
our attention. Jennifer McGown (personal com-
munication) told us of the first group. That group
consisted of a small cluster of plants in a vacant Fifth
Street lot at Nanaimo, British Columbia, approxi-
mately 35 km SE of the French Creek site. The cluster
was growing alongside a dry ditch edge amongst a
large patch of Rubus discolor. All the plants were
taller than 3 m, and all had gone to seed. The second
group was discovered near a vacant lot at Victoria,
British Columbia in circumstances similar to those of
the Nanaimo specimens (Harold Hosford, personal
communication). Specimens from these sites were
verified (T. C. Brayshaw, personal communication)
1979
and are now onfile at the British Columbia Provincial
Museum, Victoria. After the lodging of our H.
mantegazzianum species with the Provincial Muse-
um, Brayshaw informed us that another specimen was
located in the museum herbarium. That specimen,
collected in 1973 from the French Creek site, was
incorrectly labelled H. lanatum.
At the French Creek location the plants were found
in large clusters on both sides of the creek. One
collected specimen measured 395 cm from the base to
the top of the terminal umbel, with a terminal umbel
width of 70 cm. The stem was 9 cm thick at the base.
The lower 26 cm of the stem was a deep purple while
the rest of the stem was green covered with purple
spots and blotches decreasing in extent toward the
umbel. Clear hairs protruded from most of the spots.
The fruit was elliptic and ranged from 12.0 to 13.2 mm
in length. The vittae were swollen near the base and at
that point averaged slightly over | mm in width.
Further investigation revealed large numbers of H.
mantegazzianum upstream in suitable locations and
subsequently they were found growing along a
tributary of French Creek where it crosses Swayne
Road, approximately 2km S of Coombs, British
Columbia. The plants grew along the banks of the
creek, south to a point just past the intersection of the
tributary and Winchester Road. Here the local
distribution of the plant abruptly ended. French
Creek and another of its tributaries crosses Win-
chester Road a further 1500 m and 600m north,
respectively; however, an inspection of both sites did
not reveal any specimens of H. mantegazzianum.
Because of that, we feel the area of introduction of the
plant lies in the general location of the southernmost
tributary of French Creek near its crossing of
Winchester Road.
We talked with local residents, and in particular to
Frances Kroot (personal communication) who lives
near the presumed site of introduction, to try to
determine the time span H. mantegazzianum has
taken to establish itself. Apparently the plant was not
known to occur in the area in 1937; however, by 1944
it appeared sporadically where the forest edged away
from the creek, and beside bridges where openings
were created. Here the plant’s history becomes
nebulous, but by 1967, when Frances Kroot came to
the area, the plant was well established. By 1975, and
likely years before, it was known to be well estab-
lished near the mouth of French Creek, some 12 km
downstream. As Morton (1978) reports that the
principle mode of dispersal is by water, this further
supports our conclusions regarding the site of
introduction.
All the residents we talked with mentioned the
stinging effect and the blisters which occur upon
coming in contact with H. mantegazzianum, as
NOTES 83
described by Morton (1978), and all requested
information on methods of eradicating the plant. Bert
Topliffe (personal communication) described his
attempts to rid them from his yard. He made repeated
applications of the herbicide 2,4,5-T and this seemed
to be effective shortly after application. The following
year, however, they reappeared as numerous as
before. He has since found that only by continually
mowing the plants does he have any success. Topliffe
also mentioned observing a number of people each fall
collecting the large umbels. He recalled one occasion
when a couple loaded the back of their pick-up truck
with the dried stalks and umbels. This, undoubtedly,
could disperse the plant over large distances and may
account for the Nanaimo specimens.
The discovery of H. mantegazzianum on Van-
couver Island appears to warrant the concern ex-
pressed by Morton (1978). Hazards of coming into
direct contact with the plant have already been
mentioned. Perhaps more important, however, Is its
ability to become well established within a relatively
short time, likely at the expense of some of our native
species. On Vancouver Island H. mantegazzianum
established itself within 30 yr along suitable areas
of French Creek and one of its tributaries, moving a
distance of approximately 12 km within a period of
less than 40 yr. It is well established along virtually
the entire waterway from the site of introduction to
the French Creek mouth. In some locations the plant
grows 80 to 100 m away from the creek edge. The
plant’s potential for spreading even further is in-
creased through the dispersal of its seeds by humans in
their quest of dried umbels, presumably for home
decorations. Coupled with those factors are the
difficulties encountered in trying to eradicate the
plant once it is established.
As Morton (1978) points out, we need more
information as to the distribution, life history
aspects, and control methods of H. mantegazzianum
in Canada. Naturalists should also be alert to the
possibility that the piant occurs within their area and
has to date been passed off as an aberrant form of a
locally common species.
Literature Cited
Kamermans, J. K. 1977. A hairy horror in my Huron
haunts. Wood Duck 31: 49-50.
Morton, J. K. 1978. Distribution of Giant Cow Parsnip
(Heracleum mantegazzianum) in Canada. Canadian
Field-Naturalist 92: 182-185.
Taylor, R. L. and B. MacBryde. 1977. Vascular plants of
British Columbia. University of British Columbia Press,
Technical Bulletin Number 4.
Received 7 September 1978
Accepted 4 October 1978
84 THE CANADIAN FIELD-NATURALIST
Vol. 93
Nesting of Horned Puffins in British Columbia
R. WAYNE CAMPBELL,! HARRY R. CARTER,2 and SPENCER G. SEALY?
'\British Columbia Provincial Museum, Victoria, British Columbia V8V 1X4
2905 Deal Street, Victoria, British Columbia V85 5G4
3Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2
Campbell, R. Wayne, Harry R. Carter, and Spencer G. Sealy. 1979. Nesting of Horned Puffins in British Columbia.
Canadian Field-Naturalist 93(1): 84-86.
Summer observations of Horned Puffins (Fratercula corniculata) from 1972 through 1977 suggest that this species is
expanding its range along the British Columbia coast. Nesting was confirmed at one new site and probably occurred at three
others.
Key Words: Horned Puffins, Fratercula corniculata, British Columbia, breeding range.
The status of the Horned Puffin (Fratercula
corniculata) on the west coast of North America south
of its breeding range in southeastern Alaska (A.O.U.
1957) was reviewed recently by Sealy and Nelson
(1973) and Hoffman et al. (1975). These authors
considered it a late spring and early summer (and
occasional winter) visitor to this area but did not
report evidence of breeding. Forrester Island, Alaska,
was heretofore the southernmost known breeding site
in the eastern Pacific (Heath 1915: Willett 1915).
Censuses of, and other work in, seabird colonies
along the British Columbia coast from 1972 through
1977 indicate (Table 1) that the Horned Puffin is now
more widely distributed along the west coast of the
Queen Charlotte Islands and Vancouver Island than
previously known. It also has been recorded on the
central mainland coast.
Possible Nesting Sites
Marble Island (52° 12’N, 132° 39’W). This is a
fairly large, wooded island, about 150 m high, with
dense undergrowth above the precipitous cliffs. On 19
July 1977 two Horned Puffins scrambled from a
natural rock crevice on a large grassy slope among a
colony (hundreds) of Tufted Puffins (Lunda cirrhata).
Later, five Horned Puffins were seen at one time
circling off the island with Tufted Puffins. During
the half hour the Horned Puffins were watched, an
adult several times flew to, and almost landed near, its
presumed nest site. The following day two Horned
Puffins were again seen flying with Tufted Puffins off
the island.
Unnamed Islet (52°06'N, 131° 14’W). This islet, off
the northwest coast of Anthony Island, is about 30 m
high, is composed of bare rock with deep cracks and
fissures, and has grassy areas along its north tip. Six
Horned Puffins were seen flying with Tufted Puffins
close to the islet on 2 July 1977. Two days later, a
Horned Puffin flew from a crevice near the top of a
steep rocky bluff: later we discovered a recently
cracked egg in the crevice. Later the same day we
watched up to five Horned Puffins, for an hour or so,
fly very close to the islet, but none landed or entered a
natural crevice. All other possible nest sites were
inaccessible without climbing equipment. On 5
August, four Horned Puffins were flushed from rocky
vantage points above where they had been seen in early
July.
Cape St. James (51°56, 131°01’W). This landform
consists of the St. James Islands, which are saddle-
shaped, bare, and grassy with vertical cliffs about
30 m and the Kerouard Islands, which are mostly
grassy, unforested, and up to 76m high. Horned
Puffins were recorded near Cape St. James during the
summers of 1972 through 1974 and also in 1977. H D.
Fisher (letter to SGS, 26 September 1974) was not
aware “that the occurrence of Horned Puffins at the
Cape (May to August) and carrying fish at that, would
be valuable news. The common puffin in the
Kerouards is the Tufted Puffin. But among these we
always saw a few Horned Puffins. Whenever we were
enroute to and from the sea-lion rookeries, they
[Horned Puffins] seem to nest on the cliff directly
below the weather station on the northernmost
Kerouard, a very inaccessible place.”
Triangle Island (50°52’N, 129°05’W). This, the
most western of the Scott Islands, about 210 m high, is
very precipitous with no trees. Horned Puffins were
first recorded here in 1972 by C. J. Guiguet and have
been observed nearly every summer since by biologists
visiting or conducting research (Table 1; Vermeer
et al. 1976). Although positive evidence is lacking, up
to four pairs may breed on Triangle Island, all at the
western end and near “Murre” rock.
Discussion
The observations presented here and those of Sealy
and Nelson (1973) suggest that the Horned Puffin is
more common in British Columbia than previously
known. If this trend continues, opportunity will exist
NOTES 85
TABLE !|—Summer observations of Horned Puffins in British Columbia, 1972-1977
1979
Locality! Date
Queen Charlotte Islands
Cox Island (Langara I.) 31 July 1977
Darwin Sound May 1977
Marble Island 19 July 1977
Marble Island 20 July 1977
Island NW Anthony I. 2 July 1977
Island NW Anthony I. 4 July 1977
Island NW Anthony I. 5 August 1977
Island E Anthony I. 4 July 1977
Cape St. James May-August 1972
Cape St. James May-—August 1973
Cape St. James May-August 1974
Cape St. James 5 July 1975
Cape St. James 27 July 1975
Cape St. James 3 July 1977
Cape St. James 4 July 1977
Mainland Coast
Sinnett Island 27 June 1976
Joseph Island 28 June 1976
Vancouver Island
Triangle Island 1 July 1972
Triangle Island 16 June 1974
Triangle Island 2 July 1974
Triangle Island
Triangle Island
21 August 1974
June-August 1975
Triangle Island 27 July 1977
Triangle Island 28 July 1977
Triangle Island 1 August 1977
Solander Island 27 June 1975
Barrier Islands 26 June 1975
Wichaninnish Bay 20 June 1975
Number? Remarks
| Flying with Tufted Puffins
4 In a flock flying south
5 Suspected nesting
2+ Flying with Tufted Puffins
6 Suspected nesting
5 Suspected nesting
4 Suspected nesting
l Flying alone
1 or 2 With Tufted Puffins
1 or 2 With Tufted Puffins
1 or 2 With Tufted Puffins
7 With Tufted Puffins
2 With Tufted Puffins
| With Tufted Puffins
2 With Tufted Puffins
l Alone off island on water
| Flying alone
2 Flying with Tufted Puffins
I+ On water near island
2+ On water near island
| On nesting cliffs
1-6+ One carrying fish
| On rocks on island
3 On water off island
8 On rocks
3 Flying with Tufted Puffins
On water off island
! On water
‘Listed from north to south.
2All observations of birds in apparent definitive plumage.
to monitor it. Therefore, it is important to interpret
carefully our observations so that its status in British
Columbia up to 1977 is clear.
The finding of a Horned Puffin nest containing an
egg, On an unnamed islet northwest of Anthony Island
in 1977, provides positive proof of breeding. The
observations (Table 1) of birds carrying fish in their
bills is evidence short only of finding an egg or chick.
Where the above evidence is lacking, observations
must be viewed with caution. Myrberget (1959) found
that the varying proportion of non-breeding Common
Puffins (F. arctica) present in a colony may equal the
number of breeding birds there, and after 4 yr of
age many individuals prospect for and even “own”
burrows but seldom breed (Petersen 1976a). Many
more 5-yr-old birds breed but maximum reproductive
output does not occur until the birds are 10-11 yr
old. In Alaska, many occupied Horned Puffin bur-
rows during the breeding season never reach the egg
stage, possibly because their “owners” are immatures
(D. H. S. Wehle, personal communication). Such im-
matures in the Common Puffin are recognizable only
on the basis of bill shape and the number and depth of »
bill furrows (Petersen 1976b). The presence of a
Horned Puffin on a boulder or even in a burrow
therefore does not imply nesting. Also, as noted by
Lockley (1953), and confirmed by Petersen (1976a),
Common Puffins that circle the periphery of a colony
are generally immature. The breeders that are feeding
young generally fly straight out to sea on leaving the
burrows and straight back (carrying fish) when
returning. In fact, Brun (1971) considered only those
puffins carrying fish in their bills to be breeders when
he censused colonies in northern Norway. Hence the
presence of Horned Puffins of unknown age at best
provides suggestive evidence for breeding. The finding
of additional nests and additional observations of
birds carrying fish will be required to confirm that this
species is continuing to change its status in British
Columbia.
86 THE CANADIAN FIELD-NATURALIST
Another puffin, the Rhinoceros Auklet (Cero-
rhinca monocerata), is actively expanding its breeding
range (see Scott et al. 1974) and is recolonizing islands
from which it had been exterminated (see Ainley and
Lewis 1974).
Acknowledgments
We appreciate the assistance of R. Billings, T.
Carson, H D. Fisher, J. B. Foster, H. M. Garrioch,
S. M. Guiguet, M. A. Paul, M. C. Lee, M.S. Rod-
way, M. G. Shepard, and K. R. Summers with field
aspects of this work. C. J. Guiguet and J. B. Foster
kindly read the manuscript. Supporting funds were
received from the Ecological Reserve Unit, Depart-
ment of Environment, Victoria, British Columbia.
Literature Cited
Ainley, D. G. and T. J. Lewis. 1974. The history of Fara-
llon Island marine bird populations. Condor 76: 432-
446.
American Ornithologist’s Union.1957. Checklist of North
American birds. Sth edition. Lord Baltimore Press,
Baltimore, Maryland.
Brun, E. 1971. Census of puffins (Fratercula arctica) on
Nord-Fugloy, Troms. Astarte 4: 41-45.
Vol. 93
Heath, H. 1915. Birds observed on Forrester Island,
Alaska, during the summer of 1913. Condor 17: 20-41.
Hoffman, W., W.P. Elliott, and J. M. Scott. 1975. The
occurrence and status of the Horned Puffin in the western
United States. Western Birds 6: 87-94.
Lockley, R. M. 1953. Puffins. J. M. Dent, London. 186 pp.
Myrberget, S. 1959. Vekslinger 1 antall lundefgul inne ved
kolonian. Sterna 3: 239-248.
Petersen, A. 1976a. Age of first breeding in puffin, Fra-
tercula arctica L. Astarte 9: 43-50.
Petersen, A. 1976b. Size variables in puffins Fratercula
arctica from Iceland, and bill features as criteria of age.
Ornis Scandinavica 7: 185-192.
Sealy, S. G. and R. W. Nelson. 1973. The occurrence and
status of the Horned Puffin in British Columbia. Syesis
6: 51-55.
Scott, J. M., W. Hoffman, D. Ainley, and C. F. Zeille-
maker. 1974. Range expansion and activity patterns in
Rhinoceros Auklets. Western Birds 5: 13-20.
Vermeer, K., K. R. Summers, and D.S. Bingham. 1976.
Birds observed at Triangle Island, British Columbia, 1974
and 1975. Murrelet 57: 35-42.
Willett, G. 1915. Summer birds of Forrester Island, Alaska.
Auk 32: 295-305.
Received 5 April 1978
Accepted I] August 1978
News and Comment
Special Appreciation Expressed to R. Emerson Whiting
Weare particularly grateful to R. Emerson Whiting
for his careful and thorough preparation of the Index
for The Canadian Field- Naturalist from 1975 to 1978
(Volumes 89, 90, 91, and 92). His particular concern
for accuracy and his attention to details is not only
appreciated by the journal but more so by the users of
the Index. We sincerely thank Mr. Whiting for his
time and effort because the Index is such an important
component of each volume. We are pleased to
announce that W. Harvey Beck has agreed to take on
the task of indexing Volume 93.
LORRAINE C. SMITH
Editor
Second International Congress of Systematic and Evolutionary Biology
The Second International Congress of Systematic
and Evolutionary Biology (ICSEB-II) will be held at
the University of British Columbia, Vancouver,
Canada on 17-24 July 1980. The provisional list of
symposia topics include these:
1. Arctic refugia and the evolution of arctic
biota.
Origins and evolution of the north Pacific
marine biota.
Evolution of reproductive strategies.
Evolutionary epigenetics.
Evolution of community structure.
Green algae and land plant origins.
Macromolecular mechanisms in evolution.
Se stacey
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This new edition has been prepared by Warren King
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including ICBP’s National Sections and Working
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The volume covers 199 bird taxa and includes birds
that are threatened throughout their range. It does not
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87
8.
9.
10.
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Allozymes and evolution.
Coevolution and foraging strategy.
Evolution of colonizing species.
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Sessions for contributed papers and for papers in
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Those interested in receiving an information circu-
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British Columbia, 2075 Wesbrook Mall, Vancouver,
British Columbia V6T 1WS.
Resources)
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The second instalment of the World Directory of
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Now greatly enlarged the Directory is divided into
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88 THE CANADIAN FIELD-NATURALIST
Societies and Amateur Ornithologists
The following are taken from the recommendations of the
National (USA) plan for Ornithology submitted to the
National Science Foundation and the Council of the
American Ornithologists’ Union (AOU).
10. Professional societies (preferably the AOU or
an intersociety consortium) should form a committee
charged with (a) developing projects of national
ornithological importance for amateurs, (b) develop-
ing centers of responsibility for particular projects,
and (c) providing professional assistance in securing
and maintaining financial support for the projects.
11. The ornithological societies should undertake
the development of a network of involvement by local
bird clubs and nature centers. The initial step should
be the preparation of a computerized listing of all
clubs and contacts. The ultimate goal should be a
Federation of American Bird Clubs or an ATO
(American Trust for Ornithology) modelled on the
British counterpart.
12. The AOU should take the initiative in improv-
ing formal coordination in the activities of the
ornithological societies and in the work of the profes-
sion in general. An obvious option for this, perhaps
only as a developmental stage, would be the formation
of an intersociety coordinating council whose role
would be to consider or initiate long-range policy of
IUCN Views on Whale Management
The International Union for Conservation of
Nature and Natural Resources (IUCN) is committed
to the sustainable use of natural resources, including
the living resources of the seas. It believes that it is
possible to manage whaling ona sustainable basis and
is concerned, therefore, that after three years the
International Whaling Commission’s (IWC’s) New
Management Policy has not been able to achieve this
goal.
Although the IWC and its Scientific Committee
have made commendable efforts to improve man-
agement procedures, the outcome has been generally
unsatisfactory and there have been some extra-
ordinary lapses; for example, the manner in which the
quota for North Pacific Sperm Whales was sharply
raised in December last year on the basis of
questionable data and an inadequate model.
The IUCN is also deeply concerned that certain
member states so far have not met the IWC’s
requirement that full and accurate data and analyses
be furnished to the Commission. Those few states
complying find their data subjected to intense scrutiny
Vol. 93
national or general professional scope, as well as to
assist in the integration of routine activities.
13. The ornithological societies should take
greater interest in the work of ornithological centers
such as organized bird observatories (e.g., Manomet,
Point Reyes Bird Observatory) and aid in their
development and promotion.
14. The ornithological societies should establish
training programs in basic ornithology and research
techniques for amateurs. Amateur-oriented work-
shops supervised by professionals should become a
regular part of annual meetings.
15. Annual meetings of the ornithological
societies should be better structured to promote
contacts and communication between amateurs and
professionals and between junior and senior mem-
bers. Part of this goal can be reached through
Recommendation 14, but this will not be sufficient.
Explicit procedures should be devised, such as
scheduled round-table discussions, that will overcome
diffidence or reticence and automatically put ama-
teurs and professionals and junior and senior mem-
bers in personal contact in an arena of common
subject-matter interest.
From ASC Newsletter 6(5): 53-54, 1978.
(Association of Systematics Collections)
while those not complying may be awarded quotas in
line with past custom rather than with the present
conditions of the stocks. The work of the IWC is
undermined by member states when they do not
provide requested data and when they encourage
whaling activities of non-members and report less
than their total catch. IUCN suggests to the !WC that,
to end such practices and to allay suspicion, it requires
its members to provide original catch records and it
re-opens discussion of the possibility of placing truly
international observers, accredited by the IWC, to all
whaling operations.
IUCN urges the IWC and member states
— to ensure that the quotas they set this year are
conservative, particularly with respect to those
stocks about whose status there 1s considerable
doubt and those stocks which are part of multi-
species fisheries;
—to ensure that current management policy is
thoroughly revised in time for greatly improved
procedures to be applied in 1979;
—to make good their commitment to gather
1979
adequate data on whales and the ecosystems of
which they are part by implementing without
further delay the International Decade of
Cetacean Research;
— to accelerate the revision of the Convention and
ensure that its conservation provisions are
considerably strengthened and the needs of
whale conservation adequately catered for in
related instruments for the management of
marine resources.
Failure to take these measures will confirm IUCN
and the rest of the international community in their
view — expressed repeatedly since 1972 — that there
is no rational alternative to a moratorium on com-
mercial whaling.
With the support of the United Nations Environ-
ment Programme and the World Wildlife Fund,
“To Know Ourselves”
The history of Canadian science and technology 1s
still being so badly neglected by our universities that
“Canadian students know virtually nothing about
their scientific heritage,” says Thomas Symons, chair-
man of the Commission on Canadian Studies.
As a result, Symons said in a recent interview, “the
technologists and scientists of the country are often
underutilized and underestimated by their fellow
citizens. A great deal of major scientific and techno-
logical work is commissioned outside the country
because there just isn’t the realization that we have the
capability to do it here.” Symons is the author of the
Commission’s 1975 report, To Know Ourselves, one
chapter of which was devoted to science, technology,
and Canadian studies.
The Science Council of Canada is preparing an
“issues paper” on science and education in the
Canadian context. A recent Council-sponsored sem-
inar concluded that there is, at present, “no formal
requirement and little interest in putting Canadian
content in science courses.” In anticipation of a
possible full-scale study in the future, the issues paper
will attempt to define and to focus on a number of the
serious science-and-education problems highlighted
in the Symons report.
Symons said he is encouraged by the fact that halfa
dozen universities have discussed with him the possi-
bilities of either introducing or extending programs in
the history of science. A major conference on the
history of science was planned for Queen’s University
in November 1978. He also said there is some evidence
that the attitude of Canadian scientists has been
shifting in favor of the idea of Canadian content in
science teaching. This shift has, perhaps not sur-
NEWS AND COMMENT 89
IUCN has embarked on an expanded program for
marine conservation. Three of the objectives of this
program are to launch an international system of
cetacean sanctuaries, promote the conservation of the
living resources of the southern ocean, and develop
improved principles and procedures for the man-
agement of whaling.
IUCN welcomes the participation of the IWC in the
operation of the Convention on International Trade
in Endangered Species of Wild Fauna and Flora, and
hopes that through this and other joint endeavors by
concerned governments and international bodies the
current problems of the conservation of whales will be
satisfactorily resolved.
From IUCN Bulletin. New Series 9 (7/8), July/ August
1978.
prisingly, been most pronounced in the environmental
sciences.
In his report, Symons acknowledged that science is
universal and international, but he also argued
strongly that it nevertheless has a “national and
cultural dimension . . . Science is a key ingredient in
the cultural fabric of our society.” He suggests that
there are “Canadian perspectives, Canadian applica-
tions, Canadian motivations (and) Canadian ap-
proaches to science that could be described accurately
as Canadian studies.”
His report in fact contains a litany of scientific
subjects that could easily, and without rationaliza-
tion, be classified within the ambit of Canadian
studies — subjects which the Commission felt were
being woefully neglected. For example, it urged that
more attention be paid in science education to the’
implications of Canada’s weather and climate; to
marine sciences concerned with the continental shelf
and offshore conditions; to the geology and geo-
graphy of Canada’s land masses; to Canadian
mammals; and to Canada’s forests.
The report particularly emphasized the lack of
research in and about the North and the need for a
more sustained and indigenous scientific effort there.
It strongly recommended that a university of the
North, or at least its nucleus be established immed-
lately.
The Commission also argued that non-science
students in university must be given some training in
science and the history of Canadian science. This is
needed “to equip non-science students with a better
knowledge and understanding of the important part
played by science in our society . . . As matters now
90 THE CANADIAN FIELD-NATURALIST
stand, universities in Canada are producing each year
thousands of graduates whose knowledge and under-
standing of the role played by science in our total
culture is minimal to zero.”
Conversely, science students must be allowed to
take courses in the arts and humanities that will give
them a basic knowledge of the society in which they
live and the social institutions which will inevitably
affect scientific research.
Achieving this goal is obviously not an overnight
kind of problem, but Symons said that a “modest
beginning” has been made; a number of universities
have indicated to him that they are taking a “fresh
Vol. 93
look” at the problem.
On the whole, in fact, Symons conveys a generally
optimistic message. Although much remains to be
done, he feels the Commission has been successful in
prodding the scientific community to think about
these issues. He also believes that the fact that he is not
a scientist, and therefore could not be accused of self-
pleading, helped in getting a fair number of the
recommendations acted upon.
From Agenda, August, 1978.
(Bulletin of the Science Council of Canada)
Second Annual International Wildlife Film Festival
To encourage the production of high quality
wildlife films, the University of Montana Student
Chapter of the Wildlife Society sponsored the First
Annual International Wildlife Film Festival last year.
It was a great success and the chapter is again hosting
this event. A panel of highly qualified film makers and
biologists will judge films pertaining to wildlife and
present awards and recognition to the winning films.
The deadline for submission of applications and
films is 7 February 1979. All entries must have a
predominantly wildlife theme and have been pro-
duced or released in calendar year 1978. Judging will
be held on 10 and I! February and the winning films
will be shown on 3 and 4 March at the University
Center.
Information, rules of eligibility, and application
forms can be obtained by writing: Wildlife Film
Festival, Wildlife Biology Program University of
Montana Missoula, Montana 59812 (Telephone (406)
243-5272).
Book Reviews
ZOOLOGY
A Guide to Bird Finding East of the Mississippi
By Olin Sewall Pettingill, Jr., with illustrations by George
Miksch Sutton. 2nd Edition: 1977. Oxford University
Press, New York. xxvii + 689 pp. U.S. $15.95.
When I first began watching birds almost 20 years
ago, one of the first books I acquired was O. S.
Pettingill’s A Guide to Bird Finding West of the
Mississippi, published in 1953. This book and its
eastern counterpart, published in 1951, have been
constant and valued companions during my travels
throughout the United States. Now a new and
extensively revised version of the eastern guide has
been issued, and a welcome event indeed this is for
birdwatchers. Pettingill is presently revising the
western guide, with a tentative publication date set for
1980.
The format of the book is the same as in the first
edition. Following a brief introduction, it is divided
into 26 chapters, one for each American state lying
entirely east of the Mississippi River (except that
Connecticut and Rhode Island are dealt with to-
gether, and the New York City region rates its own
chapter). Canada is not covered, although several
Canadian localities near the U.S. border (e.g., Point
Pelee, Ontario and Grand Manan Island, New
Brunswick) are discussed under the nearest state.
Chapter length varies from 16 pages (Vermont) to 48
pages (Florida). Each chapter consists of two parts: a
general account of the state’s birdlife, and a list of
specific bird-finding localities, which are discussed
under the nearest sizable city or town. Each locality
account includes detailed directions for reaching the
area and a partial list of birds to be expected, with
emphasis on uncommon or spectacular species or
noteworthy bird concentrations. Both the general
state accounts and specific locality accounts, es-
pecially the latter, have been revised substantially
from the first edition. As Pettingill notes, the changes
in 25 years have been profound, many bird-finding
localities have been “developed,” and the distribution
of the birds themselves has changed in many cases.
Thus some localities have been dropped from the new
edition, but many others have been added; in
Michigan, for example, 29 localities are described
(under 18 cities and towns) in the new edition, versus
16 in the old edition. Locality accounts are generally
briefer in the new edition, but for most states, the net
result is better coverage overall.
The introductory sections in each state chapter are
especially illuminating. Pettingill begins with a
91
description of the physiographic regions of the state
and their associated vegetation and land-use types.
Then come lists of characteristic breeding species for
major habitats, with notes on those species that are
restricted to certain parts of the state. Next follows a
description of bird migration patterns (spatial and
temporal) in the state, and finally a brief account of
winter birdlife. Having thus “set the stage,” Pettingill
then fills in the details in the individual locality
accounts which follow.
The book’s index is designed for easy location of
information on a particular species. Suppose you
want to find Bachman’s Sparrows in Alabama. A
glance in the index under “Bachman’s Sparrow” tells
you that the species is mentioned in Alabama locality
accounts on pages 9 and 20; thus you need not wade
through the entire Alabama chapter.
For English names of birds, the book follows the
A.B.A. Checklist, published by the American Birding
Association in 1975. Many of these names have not
yet been adopted by the American Ornithologists’
Union, and are not yet in general use; however, as the
A.O.U. will likely adopt most of them soon, their use
by Pettingill may help to prevent the book from
becoming rapidly out-of-date. (The first edition
includes some names now long obsolete, such as
“Holboell’s Grebe” and “Duck Hawk.”)
The route directions in most cases are necessarily
brief. But I have not experienced the difficulties
encountered by one reviewer (J. A. Tucker, 1977,
Birding 9: 226-231) in following them. The birder
would certainly be advised to have a good road map:
handy, especially when trying to find localities in and
near large cities. Given the frequent changes in road
locations and numbering systems, however, directions
for reaching some areas are bound to become out-of-
date rather quickly.
In looking closely at the chapter on Mississippi (the
state I know best), I noted a couple of typographic
errors in place-names, and several errors of fact,
mostly concerning bird distribution; however, such
errors are few and of minor importance. If I do have
any serious criticism of the book, it is that, for some
states at least, the author has relied too heavily on old
information. (For instance, of 14 persons cited as
contributing information for the Alabama and Mis-
sissippi chapters, only two were not cited in the 1951
edition.) I am not suggesting that some of the areas
included are no longer good for birds—only that local
92 THE CANADIAN FIELD-NATURALIST
observers now know of better and more accessible
areas than some of those included. But I hasten toadd
that the selection of localities to include in a book like
this is difficult and arbitrary, and no such selection
will please all observers.
| have now had a chance to use the new “Pettingill”
in seven states, and I unreservedly recommend it to
anyone looking for birds in the eastern U.S. Although
there are now many good bird-finding guides for
individual states, Pettingill’s is the only one covering
the entire eastern part of the country, and | doubt that
The Birds of New Brunswick
By W. Austin Squires. 1976. New Brunswick Museum,
Saint John, New Brunswick. Second Edition. v + 220 pp.,
illus. No price given.
Ce livre est une nouvelle édition complétement
revisée de celle de 1952 épuisée depuis plusieurs
années. L’auteur y présente, en 221 pages, la situation
ornithologique du Nouveau-Brunswick. L’ouvrage
est agrémenté de dix photographies en couleur et
autant en noir et blanc; le choix est judicieux et
intéressant. J’al apprécié particulierement celles du
Martinet ramoneur, de la Sitelle a poitrine rousse, du
Grand Puffin et du Pica dos noir. La bibliographie est
complete pour la période séparant les deux éditions; il
est regrettable que l'on ne retrouve pas dans cette
nouvelle édition, toutes les références bibliographi-
ques utilisées dans la premiere. Deux index, un
regroupant les noms anglais et scientifiques suivi d'un
second pour les noms frangais, indiquent le caractere
biculturel de la province. Grosso-modo, le livre peut
étre divisé en deux sections: l'une traite de différents
aspects de lornithologie dans la province et la
seconde, constituant la majeure partie de l’ouvrage,
est une liste annotée des oiseaux.
Dans la premiére section, l’auteur, en plus d’in-
troduire la liste annotée, fournit quelques courts
chapitres sur des sujets aussi divers que Ihistoire, la
biologie, l’écologie et la protection des oiseaux tout en
limitant sa discussion a la province. Le chapitre sur
Phistoire de lornithologie est lélément le plus
original. L’auteur cerne trés bien le développement de
lornithologie de 1613 a 1952; cependant, la période de
1952 a nos jours, période ot lornithologie-amateur
contribue de fagon importante au développement des
connaissances sur la distribution des oiseaux, est
déficiente. Ainsi, 1] aurait été intéressant d’avoir des
renseignements sur les groupes d’amateurs et de
professionnels oeuvrant dans le domaine. A certains
endroits de ce chapitre, 11 est difficile de distinguer
entre les dates de visites des personnages et celles du
Vol. 93
his approach will ever be significantly improved upon.
Pettingill is to be congratulated on producing a well
done and much needed revision of a book which is
already considered a classic. My only lament is —
when will someone write a bird-finding guide of
similar comprehensiveness and authority for Canada?
WAYNE C. WEBER
Department of Biological Sciences, Mississippi State Uni-
versity, Mississippi State, Mississippi 39762
systeme de références bibliographiques; une typo-
graphie différente faciliterait la consultation. Lors de
la révision de l’édition originale, on aurait dQ apporter
une attention particuliére a la conversion des mesures
dans le systéme international (SI). Une carte de la
province représente cing régions topo-climatiques, les
comtés provinciaux et quelques repéres géographi-
ques importants. Un lecteur, non-résident du Nou-
veau-Brunswick, apprécierait une carte avec une
toponymie plus complete en relation avec la liste
annotée. Ainsi, je n’ai pu localiser Pointe Lépreau,
localité fréquemment mentionnée dans louvrage; de
méme que l’ile Nantucket, homonyme de la célébre ile
au large du Massachusetts. Les autres chapitres de
cette section sont intéressants: l’introduction et les
chapitres sur la migration, la collection et la ter-
minologie de la liste annotée doivent étre consultés
avant la lecture de la seconde partie de Pouvrage.
Cette seconde section fournit une liste annotée de
345 espéces d’oiseaux. Dans la présentation de ce
genre de listes, deux approches peuvent étre utilisées;
Pune consiste a décrire l’espéce, son habitat et
quelques traits de sa biologie, suivi de son statut
provincial ou: régional. C’est approche que lon
retrouve dans des ouvrages concernant d’autres
provinces canadiennes (Salt, W.R. et J. R. Salt,
1976, The birds of Alberta; Tufts, R. W., 1961, The
birds of Nova Scotia). L’autre présentation consiste a
se limiter au statut de l’espéce, en le précisant par une
sélection de mentions. C’est cette derni¢re approche
qu utilise Squires et, a mon avis c’est celle quiatteint le
mieux les objectifs que ce genre de livre devrait avoir.
Dans la présentation de chaque espéce, les noms
anglais, frangais et scientifiques sont fournis suivis
dun bref résumé du statut et de l’'abondance selon des
termes préalablement définis. Pour la majorité des
espéces, une bréve description de lhabitat et une
sélection de mentions typiques viennent préciser
Vabondance. Deux autres paragraphes décrivent
1979
respectivement la distribution générale et provinciale.
Enfin, les dates de ponte pour les nicheurs et la période
de résidence de lespéce au Nouveau-Brunswick
terminent le traitement de chaque espeéce. Ce genre de
présentation agrémente la consultation de cette liste.
L’auteur a fouillé en profondeur la littérature con-
cernant les mentions pouvant se rapporter au ter-
ritoire considéré. Cependant, le traitement d’espeéces
marginales, quoique précis, souffre d’une déficience
bibliographique. Ainsi, auteur aurait pu faire le
rapprochement des invasions de Vanneau huppé en
1927 et 1966 avec celles des provinces voisines; les
mentions de Fauvettes azurées pour le Québec sont
basées sur des spécimens (Ouellet, H., 1967, The
distribution of the Cerulean Warbler in the province
of Quebec, Canada, Auk 84: 272-274) et non
uniquement sur des observations visuelles. Le traite-
ment des espéces occasionnelles et accidentelles donne
BOOK REVIEWS 93
souvent prise a la critique dans ce genre d’ouvrage;
cependant, l’auteur les aborde cas par cas avec un
jugement critique.
Cette seconde €dition est présentée sur papier de
qualité avec une typographie claire et dégagée. Je n’y
ai relevé que quelques erreurs typographiques; la seule
importante est la référence bibliographique de Tufts
(1961) concernant le spécimen d’Ibis blanc. Enfin, je
consideére que cet ouvrage est indispensable a ceux qui
sintéressent a la distribution des oiseaux dans le nord-
est de Amérique du Nord et particuliérement au
Canada.
PIERRE LAPORTE
Service canadien de la faune, Région du Québec, 2700
boulevard Laurier, CP 10100, Edifice “A”, Ste-Foy,
Québec GIV 4H5
Mountain Monarchs: wild sheep and goats of the Himalaya
By George B. Schaller. 1977. University of Chicago Press,
Chicago. 425 pp., illus. US$25.
According to the author, Mountain Monarchs is“a
companion volume to The Deer and the Tiger in
which I treated the ecology and behavior of some large
mammals of peninsular India.” For Mountain Mon-
archs, Schaller restricts himself, geographically, “to
the Great Himalaya of Nepal and India, and to the
ranges of Pakistan,” and to the subfamily Caprinae.
“Rather than concentrating on one kind of animal
in one locality for years, in the accepted manner of
current ungulate studies, | moved from area to area,
spending one month with one species, then the next
one with a different species.” This approach, coupled
with a fairly extensive literature review, allows
Schaller to discuss a variety of caprine species from
a comparative standpoint. It is this comparative
treatment of the species that makes the book
enjoyable reading.
In reviewing the Caprinae, Schaller comments on
taxonomy, distribution, physical attributes, popula-
tion dynamics, herd dynamics, predators, aggressive
behavior, courtship behavior, mother—young rela-
tions, and social behavior. Chapters I found particu-
larly interesting were those dealing with “Herd
Dynamics,” “Aggressive Behavior,” and “Courtship.”
“The basic unit in caprid society consists of a female
with her offspring. All other social systems represent
permutations of this basic theme.” “Most Caprinae
herds are open, in that animals join and part, often
when population density is great and not as often
when it is sparse.” Neither of these comments alone is
particularly profound; however, they are significant in
that they provide confirmation of caprine behavior
not previously studied.
Chapter 9, dealing with “Aggressive Behavior,” is
probably the strongest and most thorough chapter in
the book. It represents a significant review of
aggressive behavior and provides comparative data
for species inhabiting the Himalayan region.
Chapter 10, dealing with “Courtship Behavior,” is
not as well organized, but is equally thorough. By
comparison chapter I1, “Mother—Young Relations,”
is far from thorough. It is probably the weakest
section in Mountain Monarchs.
The last chapter is somewhat of asummary chapter. °
“When one looks for correlations two environmental
variables obtrude: (1) species live longer in a habitat
with a predictable food supply, and (2) species
dwelling on and around cliffs live longer than those on
flat to rolling terrain.” Schaller’s work is intended to
provide some basic data on the status of unknown big-
game populations in Asia. It is his hope that
conservation efforts to preserve these species will be
soon forthcoming. “In writing about this vanishing
mountain fauna I feel a special urgency. All too often
in history the last of a species has disappeared into the
belly of a hungry hunter, its epitaph a belch.”
PETER CROSKERY
Ontario Ministry of Natural Resources, Ignace, Ontario
POT ITO
94 THE CANADIAN FIELD-NATURALIST
Vol. 93
Précis de Zoologie: Vertébrés. 3 — Reproduction, biologie, évolution et systématique. Oiseaux et
mammifeéres
Par P. P. Grassé. 1977. Masson, Paris. 2e édition. 395 pp.
$35.00.
Ce volume fait partiellement suite au Précis de
Zoologie, Vertébrés, de la méme collection, qui
réunissait Anatomie comparée, Biologie et Systéma-
tique. Afin de présenter les faits avec plus d’ampleur et
d’offrir un apercu plus fidéle de l’état actuel de nos
connaissances, l’ouvrage a été divisé en trois volumes
indépendants et sensiblement augmentes.
C’est un livre de format pratique (16 x 24 x 2cm; |
kg), trés bien relié et d’une solidité qui devrait résister
a l'usage le plus soutenu. En feuilletant le livre, j'ai été
favorablement impressionné par la qualité du papier
glacé et la clarté du texte imprimé. II y a plusieurs
dessins a l’encre de chine, quelques photographies
noir et blanc un peu floues ainsi qu'un croquis en
couleur qui n’apporte toutefois rien a la compreé-
hension du texte.
Le présent livre traite de la reproduction, de
Pévolution, des particularités physiologiques, de
léthologie et de la systématique des classes des
oiseaux et des mammiferes. La partie qui concerne les
oiseaux occupe le tiers du livre, celle qui traite des
mammifeéres occupe le reste. On y décrit en détail les
particularités anatomiques de ces animaux. Par
exemple, l’auteur fait une excellente description des
adaptations morphologiques qui permettent aux
oiseaux de voler. Une section moins importante est
consacrée a la systématique. On y présente les
Ways of the Six-Footed
By Anna Botsford Comstock. 1977. Cornell University
Press. Ithaca. x11 + 152 pp. U.S. $5.95.
This reissue of the 1903 edition is subtitled on the
dust jacket: “A delightful introduction to the world of
insects” — so it is! The author’s poetic yet intricate
description of the lives of a few common insect species
provides the reader with new insight into the lives of
insects. The ten chapters cover such subjects as sound
communication, the Maple-leaf Cutter, mimicry,
socialism, the Mud Dauber and Mason Wasp, the
Carpenter Bee, the Basswood Leaf-roller, the Lace-
wing, the Seine Maker (Hydropsyche), and the
Seventeen Year Cicada. There are 47 illustrations,
many of them the author's, for, as stated by Edward
H. Smith in the new foreword, “she became one of the
foremost wood engravers of her time... .”
Not only did she illustrate books for her husband,
caractéristiques anatomiques ainsi que quelques
aspects des moeurs des principaux genres.
Malheureusement, les descriptions sont peu détaillées
et le lecteur ne peut pas tellement simaginer lallure
des animaux décrits. Enfin, une section est consacrée
au comportement. Cette partie de louvrage est tres
décevante; le choix des sujets a été fait de fagon
arbitraire et chacun des aspects comportementaux
s€lectionnés est traité sommairement.
Le volume contient aussi quelques hors-textes: un
avant-propos, une table des mati€res et un index
alphabétique des sujets. Cet index est trés bien fait et
permet de trouver rapidement les informations
désirées. L’auteur n’a cependant pas incorporé de
bibliographie a son ouvrage, ce qui constitue une
lacune importante puisque ce traité ne couvre que
superficiellement la majorité des sujets.
En conclusion, malgré mes remarques critiques, je
crois quils’agit tout de méme d’un ouvrage didactique
de qualité, du moins ence quiconcerne l’enseignement
de la morphologie et de la systématique des oiseaux et
des mammiféres. Quant aux moeurs de ces animaux,
le lecteur y trouvera une foule dinformations qui
Yaideront a s’en faire une bonne idée.
JEAN Luc DESGRANGES
Service canadien de la faune, Région du Québec, 2700
boulevard Laurier, C.P. 10100, Ste-Foy, Québec G1V 4H5
John Henry Comstock, but co-authored several, as
well as publishing some of her own. Her purpose in
writing the stories in the present volume was to
illustrate the truth that “wherever there is life there are
problems confronting it; and that the way of solving
these problems has been the way to success in the
evolution of a species.”
The author has dedicated her book “to all those
who have been my good comrades and fellow loiterers
in nature’s byways” and it is to these people, or, in
other words, all naturalists, that | would recommend
this book.
WILLIAM B. PRESTON
Manitoba Museum of Man and Nature, 190 Rupert Avenue,
Winnipeg, Manitoba R3B 0N2
1979
BOOK REVIEWS 95
Birds of Southeastern Michigan and Southwestern Ontario
By Alice H. Kelley. 1978. Cranbrook Institute of Science,
Bloomfield Hills, Michigan. vi + 99 pp., illus. Paperback
US $2.95.
Alice Kelley’s book is the latest regionally specific
reference on birds in Canada and adjacent United
States. Records of birds are taken from the area
bounded by Kettle Point on the southeast end of Lake
Huron, Rondeau Provincial Park, Toledo, and Flint.
The author includes a brief four-page description of
the topography, drainage, soils, status of forest and
marshland, and areas of particular ecological interest.
Although short on illustrations, consisting solely of
a map of the region surveyed, the book is long on
detailed bird records, providing data on 338 species
for the period 1945 to 1974. Each species is listed with
abundance rating, residency, record arrival and
departure dates, average arrival and departure dates,
exceptional sightings and abundance records, breed-
ing status, and locations and sites historically favored
to viewing the birds.
One of the main objectives of Alice Kelley in writing
the book is to document the historic changes in bird
occurrence in order to monitor effects of rapid
environmental alterations. Therefore, the book is
written primarily for the seasoned birder/ ornitholo-
gist who has advanced beyond basic identification to
the point of determining geographic or ecological
significance of particular bird sightings at specific
locations.
Watching Sea Birds
By Richard Perry. 1975. Taplinger Publishing Company,
New York (Canadian distributer Burns and MacEachern,
Toronto). 230 pp., illus. $13.25.
There were few books written on the natural history
of seabirds of the British Isles in the 1940s that were
better, or contributed more to the subject area, than
Richard Perry’s Lundy, Isle of Puffins (1940) and
Shetland Sanctuary (1948). The information con-
tained in these publications was extensive and
important at the time; included were precise docu-
mentations of the breeding activities of colonial-
nesting seabirds at two major sites and numerous
unique interpretations of their biological significance.
Thus, the republication of these long out-of-print
classics under the title Watching Sea Birds is a most
welcome event and will allow general readers and
students of seabird biology an opportunity to gain
easy access to otherwise scarce sources of very useful
material.
Two disappointing features of the book are these:
(1) the I5- x 23-cm format which is unsuitable for
most pocket sizes and therefore relatively incon-
venient to carry in the field as a complement to
most field identification guides;
(2) as the author suggests, the general qualitative
statements regarding abundance and status have
not been rigidly defined in quantitative terms. In
1954, the New York State Federation of Bird
Clubs outlined numerical standards for quali-
tative terminology, and these could have been
incorporated into the report as extensive quanti-
tative data were available. This arbitrary system
of abundance rating would invalidate most
comparisons with those reported in other regional
summary accounts using the established stan-
dards (e.g., History of the birds of Kingston,
Ontario by Helen Quilliam).
Disregarding these minor detractions, Birds of
southeastern Michigan and southwestern Ontario is a
first-rate, comprehensive, regional documentation of
birds recommended for those wishing to apply greater
significance to their bird sightings and possibly
contribute meaningful records for that region.
BRENT BEAM
Beak Consultants Limited, 6870 Goreway Drive, Missis-
sauga, Ontario L4V I1L9
The volume under review is a straightforward
reprint of the earlier works containing the identical |
text with only minor editorial changes, usually most
evident by the removal of out-dated or technically
unacceptable sentences and paragraphs. The first half
presents the main chapters from the Lundy book on
spring occupancy, Atlantic Puffins, Black-legged
Kittiwakes, Razorbills, and Common Murres, while
the remainder comprises those from the Shetland
book on Great Skuas, Arctic Skuas, Northern
Gannets, and northern Common Murres. These
accounts are the results of intensive studies conducted
by the author from 20 March to 7 August 1939 on
Lundy Isle in southwestern England and on the Isle of
Noss, in the Shetland Islands, from 2 April to 18
September 1945. They provide a detailed and accurate
description of the breeding biology of the various
species, with particular emphasis placed on behavior,
and most of the material covered is as significant and
96 THE CANADIAN FIELD-NATURALIST
informative today as it was when first produced.
Together, they present a vivid and authoritative
account of a community of seabirds made by anastute
field observer and go far in capturing and translating
the very special atmosphere associated with the study
of colonies of seabirds and their islands.
There are, however, a few things wrong with this
new version; for the most part these are errors in
editorial judgment. The flagrant attempt by the
author and/or publisher to disguise the reprint as a
“new” book by not pointing out the existence of the
original works or even the years in which the studies
themselves were made (nowhere are the years given)
was unwise and totally unwarranted. Although this
decision was undoubtedly based on marketing con-
siderations, it is difficult to understand how it came
about; if anything, the potential for sales should have
been enhanced by announcing the volume as a reprint
of the earlier books, especially since copies of the
originals are difficult and costly to obtain. The
omission of all the photographs used to illustrate the
earlier books is unfortunate, especially those in the
Lundy book, which were carefully integrated with the
text to show precise features of certain breeding
activities. The substitute black-and-white ink sketches
of birds in flight and on land by Richard Richardson
do not fill this gap and are, in general, uninspiring and
often inaccurate, adding very little to the book. Also,
the decision not to reprint the summaries of the
breeding schedules for individual species or the
appendices (these contain statistical data on nesting
and timing, etc.) is hard to accept. A judicious
selection for inclusion should have been possible and
Bird Hazards to Aircraft: problem and prevention
By Hans Blokpoel. 1977. Clarke Irwin, Toronto, in associ-
ation with the Canadian Wildlife Service, Evironment
Canada and the Publishing Centre, Department of Supply
and Services, Ottawa. 236 pp., illus. $9.50.
That both birds and aircraft may helplessly tumble
to earth after colliding in mid-air has been known
since 1912 when the first such fatality occurred. Since
that time there have been at least 77 reported bird-
aircraft crashes with over 100 lives lost. The total
financial loss resulting from these crashes is in the
order of $100,000,000.
In spite of these staggering financial and personal
losses, and the certainty of future bird-aircraft crashes
to feed the fears of the world’s millions of flight
phobic individuals (30% of the flying population is
flight phobic for one reason or another), there has
never been a comprehensive guide or manual on the
Vol. 93
would have ensured a still wider audience by
increasing the value of the new version to both the
seabird specialist and serious naturalist. As it now
stands, these readers must of necessity go back to the
originals for this information.
The major shortcoming, however, is the absence of
an epilogue by the author to relay his assessment and
personal feelings to the reader concerning the signifi-
cance and implications of the large reductions in alcid
populations that have taken place at his study colonies
over the last 30 years. For example, the total number
of puffins, razorbills, and murres at Lundy has
decreased from about 33,000 breeding pairs in 1939
(from author’s census) to less than 2300 pairs at the
present time, with the puffin population virtually
extinct (about 40 pairs in 1969-1970). Perry has
unfortunately lost a great opportunity to add to the
biological record of the populations that he first
identified and so carefully appraised. This oversight
can only be regretted by those of us concerned about
the future welfare of this most threatened group of
specialized seabirds and of the richness and diversity
of marine ecosystems.
Overall, this new book will serve as a reliable and
accurate source of the descriptions of seabirds and
their breeding activities, and it will be found to be
informative and useful to the general reader.
D. N. NETTLESHIP
Seabird Research Unit, Canadian Wildlife Service, Bedford
Institute of Oceanography, P.O. Box 1006, Dartmouth,
Nova Scotia B2Y 4A2
of bird/aircraft collision
problems of bird hazards to aircraft. Recognizing
this fact and drawing on first-hand experience
gathered in more than 10 years of scientific study on
two continents, Hans Blokpoel presents us with a
volume designed to “. . . deal with all aspects of the
problem.” This may sound like an ambitious goal but
readers will find that Blokpoel’s objective is met very
methodically in Bird Hazards to Aircraft by (1)
examining the biology of birds as it applies to the
problem; (2) examining bird strike statistics to
determine the characteristics of the problem; (3)
examining engine designs and airports and their
surroundings to see how present and future problems
may be eliminated or alleviated; and (4) producing
references and organizations for further consider-
ation. Bird Hazards to Aircraft is written and
intended for those directly concerned with the
IQ7Y
problem, regardless of their speciality. It is very
readable.
Bird strikes generally occur to the engine of an
airplane, at night (based on strike rate), during the
spring or autumn migration, at heights under 3000
feet, during take off or landing (1.e., near airports), at
aircraft speeds of 80-160 knots and, as in the 1912
crash, the species most often involved is a gull. Birds,
however, have struck nearly all leading edges of
aircraft, at any time of the day and year, at heights of
up to 23000 feet, during all flight phases, at speeds of
up to 260 knots, and strikes have involved birds as
small as a Ruby-crowned Kinglet or as large as a
Sandhill Crane.
From here Blokpoel discusses ways of bird-
proofing aircraft and on-board means of diverting
them from the path of aircraft.
Another approach to alleviating the bird hazard to
aircraft is to reduce the attraction provided by
airports. The most common attractants at airports are
food and a safe place to nest or rest. Proper habitat
manipulation can reduce both of these. Blokpoel says,
“The airport should be made as much as possible into
a monoculture, thus supporting only a few bird
species against which effective scaring techniques can
be developed.”
Away from airports the main source of bird
problems are from local and migratory flights. It is
often possible to shift local bird flights by removing
the birds’ food sources or altering their roosting sites.
Bears — their biology and management
Edited by M.R. Pelton, J. W. Lentfer, and G. E. Folk.
1976. IUCN Publication, New Series Number 40. Mor-
ges, Switzerland. 467 pp. US $12.
Bears — their biology and management is a
selection of papers from the third International
Conference on Bear Research and Management. The
text contains 45 papers with the papers grouped into 5
sections. The 5 sections cover bear behavior, bears in
national parks, management of bears and techniques
of management, status of bears, and biology of bears.
The authors of the presented papers are international
but approach common themes and problems.
The eight papers included in section | are all the
result of observational studies on bear behavior.
Bacon and Burghardt’s paper, “Learning and Color
Discrimination in the American Black Bear,” points
out that “black bears appear to use their eyesight
during ingestive behaviors much more than pre-
viously supposed.” Luque and Stokes’ paper, “Fishing
Behavior of Alaska Brown Bears,” examines “brown
BOOK REVIEWS 97
Migratory movements of birds are much more fixed
and Blokpoel discusses bird forecasting systems that
have been devised to cope with large numbers of
migrating geese and swans.
The concluding chapter and bibliography are
excellent reference material. Listed are committees
and individual names and addresses from 33 countries
where the bird strike problem is being studied. The
bibliography contains over 400 references and cita-
tions to other bird hazard/radar ornithology bibli-
ographies.
The shortcomings of the book are few and only ofa
minor nature. I found that Blokpoel’s occasional
anthropomorphic reference made the book more
readable; others may not agree. Some pre-1973
species spellings are still employed (e.g., Widgeon).
The occasional technique is described without a
reference given (e.g., p. 103). The Appendix, Table I-
3, on ground speeds of birds could obviously have
been more complete (no gull species are given),
especially where the original covered several journal
pages. Those few shortcomings, however, are ex-
tremely minor and this excellent book should appeal to
(and be required reading for) all those, “. . . directly
”
involved in or concerned with bird strikes .. .”.
D. V. WESELOH
Canadian Wildlife Service, Canada Centre for Inland
Waters, P.O. Box 5050, Burlington, Ontario L7R 4A6.
bear fishing behavior, its development, and _ its
relation to environmental and social factors.” Other
papers of this section deal with social behavior, —
breeding behavior, and threat behavior.
Section 2 contains seven papers, all concerned with
bears in national parks. Papers of this section discuss
people--bear conflicts, movement patterns of bears,
territoriality, and bears and garbage. Beeman and
Pelton’s paper, “Homing of Black Bears in the Great
Smoky Mountains National Park,” demonstrates that
“the probability of a bear becoming a nuisance other
than in its established home range is quite low.”
Although the papers of section 2 have strong
management overtones, it is section 3 which contains
papers classed as those dealing with management.
This section contains 10 papers all of which are
interesting. Two of the papers report on bear
population control. Greer’s paper, “Managing Mon-
tana’s Grizzlies for the Grizzlies,’ suggests that
hunting is not a big factor affecting grizzly popula-
98 THE CANADIAN FIELD-NATURALIST
tions, but were hunting eliminated the grizzly popu-
lation could become a problem. Kemp’s paper,
“The Dynamics and Regulation of Black Bear Ursus
americanus Populations in Northern Alberta,” dem-
onstrates that the adult males were affecting “a
regulatory influence on the bear population.”
Section 4, the Status of bears, contains 10 papers,
mainly Eurasian in content and concerned with
declining brown bear populations. Typical of papers
contained in this section is Elgmork’s paper, “A
Remnant Brown Bear Population in Southern Nor-
way and Problems of its Conservation.” This paper
points to one of the problems. “The most reasonable
explanation for the relatively rapid decline starting in
the second half of the 1950s is therefore not hunting
and insufficient reproduction but the deterioration of
the habitat caused by increasing human activity.”
Vol. 93
The biology of bears is the topic of the 10 papers
contained in section 5. Some of these papers are
strongly physiological in slant. Rogers and Rogers’
paper, “Parasites of Bears: A Review,” I found to be
the most interesting paper of this section.
On a general level, Bears — Their biology and
management contains a wealth of information. Those
persons familiar with the 28 papers contained in the
1970 proceedings (IUCN Publication New Series
Number 23) will note that these 1974 proceedings
appear as a continuation of the earlier proceedings.
PETER CROSKERY
Ontario Ministry of Natural Resources, Ignace, Ontario
POT 1TO
The Earthworms (Lumbricidae and Sparganophilidae) of Ontario
By John W. Reynolds. 1977. Life Sciences Miscellaneous
Publications. Royal Ontario Museum, Toronto. 1x + 141
pp. $8.00.
It is with genuine pleasure that I welcome this guide
to some of the lower forms of life, and at a reasonable
price. It is regrettable that the title does not include all
of Canada; however, the book does purport to be
useful in southeastern Canada and the adjacent
United States. I suspect that with care this publica-
tion would be useful in western Canada as well, forall
but one of the nineteen species covered are intro-
duced species. Further, fourteen of these species have
been recorded from British Columbia. John Reynolds
has authored numerous technical and popular articles
on earthworms in a number of publications.
The foreword (from which I quote later) is by lanR.
Ball, Assistant Curator of Invertebrate Zoology,
Royal Ontario Museum. Included in the book are an
introduction to the general biology of earthworms,
methods of study, an illustrated glossary, a key to
species, species accounts, and a discussion of distribu-
tion and ecology of earthworms. The appendix
includes a provincial description. The book is well
documented, as indicated by fifteen pages of refer-
ences. I have followed the author's instructions on the
preservation of earthworms and have found the key
workable.
Several pages are devoted to each species account,
which is illustrated with line drawings and a map of
the distribution in Ontario. For each species a
synonymy, diagnosis, biology, range (in the world),
North American distribution, and Ontario distribu-
tion are given. Common names are given in Frenchas
well as English.
I do not hesitate to recommend this book to anyone
with an interest in the lower forms of life, especially to
he who “finds his happiness unearthing worms.”
WILLIAM B. PRESTON
Manitoba Museum of Man and Nature, 190 Rupert Avenue,
Winnipeg, Manitoba R3B ON2.
The Passenger Pigeon — its natural history and extinction
By A. W. Schorger. 1955 (reprinted 1973). University of
Oklahoma Press, Norman (Canadian distributor Burns
and MacEachern, Toronto). xii + 424 pp. $7.50.
The University of Oklahoma Press has done the
biological community a great service in reproducing
this book. The Passenger Pigeon was an incredibly
abundant species, perhaps the most abundant bird
that ever lived. Its extinction is the classic disaster
story of ecological and conservation literature.
Any species as abundant as the Passenger Pigeon
(Ectopistes migratorius) is bound to invoke all sorts of
legends. The decline of the Passenger Pigeon was close
1979
to the point of no return when scientific investiga-
tions of bird behavior and ecology were in their
infancy, and most ornithological journals were just
beginning or not yet in existence. Thus, most
information on the species was a jumble of fact and
fiction scattered through old rare books, newspapers,
and obscure journals. Schorger did an excellent job of
compiling all this scattered data, and then sifting fact
from myth. Myths are not simply dismissed, but
usually explained as misinterpretation of facts: For
example a habit of alighting on the ends of limbs of
oak and beech to eat loose nuts that had not yet fallen
caused the birds to winnow their wings back and forth
to maintain balance. This resulted in a legend that
they detached the nuts with blows of their wings.
The first chapter is appropriately devoted to early
accounts. Seekers of Canadian content will be
delighted to learn that the first known record of this
bird comes from Prince Edward Island where Cartier
saw an “infinite number” on | July 1574. The fifteen
remaining chapters document behavior, food, use by
humans, anatomy, distribution, and extinction. The
last of these chapters is a critique of various drawings
and paintings of the birds, based primarily on
anatomical facts.
In general, the book is well written, easy to read,
and remarkably free of technical errors. The caption
and legend for Figure | lack any reference to the large
nesting area shown on the map. The lengthy original
quotes will be somewhat tedious to some readers, but
a delight to others. Those of us interested in the
history of science will find the 1685 quote on the
physiology of “pigeon milk” in Columba livia fascin-
ating, but it does seem a little out of place in this book.
That the Passenger Pigeon existed in enormous
The Bird Finder’s 3-Year Note Book
By Paul S. Eriksson. 1976. Paul S. Eriksson Inc., New
York. pages unnumbered. US $7.95.
This 6- x 9-inch spiral-bound notebook is billed
as a natural companion to a field guide for anyone
wanting to maintain a permanent record of observa-
tions. Each day of the year is given one page. On this
page, space is allotted for permanently recording
observations for three years. The book provides an
organized and orderly way to record year-to-year
comparisons of migratory dates, behavioral charac-
teristics, unusual observations, identification details,
and other important notes that often become mis-
placed after the return from a field trip. Since the
maintenance of field notes can be extremely im-
BOOK REVIEWS 99
numbers cannot be doubted, but the author’s unhesi-
tating acceptance that this was the most abundant
bird ever known ignores similar estimates for the Red-
billed Dioch ( Quelea quelea) of Africa. The use of old
and obscure sources with old place names resulted in
some errors in distribution, at least in the prairie
provinces. These have been corrected by C.S.
Houston (Blue Jay (1972) 30: 221-222). One of the
most useful features of the book for the serious
biologist is the detailed documentation of sources.
Some of these lack full publishing details.
To these minor criticisms, | would add only that the
book is now over 20 years old. Serious researchers will
need to consult recent literature for an update of
information. Three more recent Canadian references
that come to mind are those by Steele (Canadian
Field-Naturalist (1967) 81: 172-174) for Ontario,
Houston (Blue Jay (1972) 30: 77-83) for Manitoba
and Saskatchewan, and Smith and Kidd (Canadian
Field- Naturalist (1971) 85: 259) for Alberta.
Schorger’s book is an excellent compendium of
information that would not have been available
without his decades of hard work. I recommend it
highly to all ornithologists. To conservationists it will
serve as an excellent example of the susceptibility of
nature to human thoughtlessness. The fact that there
exists only one known specimen of a nestling of what
may well have been the most abundant bird ever
known is shocking testimony to the lack of foresight
by man in regard to this incredible bird.
MARTIN K. MCNICHOLL
Beak Consultants Ltd., 3530 11 A St. N.E., Calgary, Alberta
T2E 6M7
portant in future years and is often ignored by even the
keenest field observer, any item that will encourage an
observer to record his many worthwhile observations
is important. Remsen (1977, On taking field notes,
American Birds 31: 946-953) recently presented a
thorough and enlightening discussion on why one
should take field notes, how they should be written,
and how they can be used.
At the bottom of each page is the inclusion of a
quotation by a famous writer, an observation by
Eriksson, or space for additional notes. Among the
writers quoted are Peterson, Burroughs, Sutton, and
Pettingill. A Peterson quote summarizes the im-
portance of recording detailed field observations, “If
exact numbers of each species are kept, a year-to-year
100
comparison gives a hint of increases or declines.
Redstart — common does not mean much, but
redstart — 58 does.”
A special feature at the back of the book isa 10-page
life list of more than 700 species listed alphabetically
by their common and scientific names with space
provided after each name for writing. Although this
feature is an excellent idea, the list contains several
errors that may confuse the user. Names are based
upon the latest update (32nd supplement) of the
A.O.U. Check-list, but usage is not consistent. Among
the confusing names discovered in this section are
Trudeau’s Tern, a South American species (type
specimen taken by Audubon in New Jersey); White
Ibis, also listed as Spanish Curlew; Common Golden-
eye, also listed as Whistling Duck; scientific name for
BOTANY
THE CANADIAN FIELD-NATURALIST
Vol. 93
Snow Goose given as Chen caerulescens hyperborea;
Dipper listed under Ouzel, and Gray Hawk also listed
as Mexican Goshawk. The treatment of the junco
group is also very confusing. Misspellings spotted in
the life list or diary sections are Gadwal for Gadwall,
Anus for Anas, Widgeon for Wigeon, and albifrans
for albifrons.
In summary, this type of diary may benefit many of
us who find it difficult “to start from scratch” and to
continue the endeavor of maintaining an organized
record of field observations after a time.
NOEL J. CUTRIGHT
Wisconsin Electric Power Company, Milwaukee, Wis-
consin 5320]
The Rare Vascular Plants of Ontario/Les Plantes Vasculaires Rares de Ontario
By George W. Argus and David J. White. 1977. Syllogeus
Series Number 14. National Museum of Natural Sciences,
Botanv Division, Ottawa. 64 pp. English + 67 pp. French.
Paper free.
The systematic conservation of our native flora
requires current, accurate information concerning the
distribution, abundance, and autecology of each
taxon. Accordingly, taxa that are found to be rare
and/or threatened can receive priority for conserva-
tion. Following this strategy, the Systematics and
Phytogeography Section of the Canadian Botanical
Association established a Rare and Endangered
Plants Committee in 1973 to enumerate the rare and
potentially endangered native plants in Canada. In
concert with these efforts, the Botany Division of the
National Museum of Natural Sciences initiated a
project to compile a data base on rare and endangered
Canadian plants. In the authors words, The Rare
Vascular Plants of Ontario is “the first in a proposed
series dealing with the rare vascular plants of the
provinces and territories of Canada.” (At the time of
review similar lists in the Syllogeus Series were also
available for Alberta and Nova Scotia.)
This publication presents a neatly abbreviated
synthesis of information about the rare, native
vascular plants of Ontario drawn from many sources:
plant distribution maps, literature, IBP/CT check-
sheets, the examination of collections in the herbaria
at the National Museum of Natural Sciences (CAN)
and the Department of Agriculture (DAO), and the
considerable input of botanists and naturalists knowl-
edgable about the flora of Ontario, who were
requested to comment on preliminary lists compiled
from the preceding sources.
Specifically, the contents of this publication include
an introductory section, an annotated list of rare
plants, a bibliography, and three appendices. The
introductory section briefly explains the history of the
project, and gives definitions of rare, endangered, and
threatened plants, the organization of the list, future
plans for the continuation of this work, potential
application of the list for the conservation of the
Ontario flora, and notes about the distribution
patterns of rare plants in Ontario. The considerable
acknowledgments support the author's tenet “that no
one person or group has sufficient knowledge of the
flora of Canada to make a project of this magnitude a
success without the support of all botanists and
naturalists.” The present work is a tribute to the
authors’ ability to marshal these human resources.
Although the authors supply definitions for ‘rare,
‘endangered,’ and ‘threatened, the present work, as
suggested by the title, is clearly an enumeration of
native vascular plants considered by the authors to be
rare. ‘Plant’ is defined “to indicate collectively all the
individuals referable to a plant name” including
species, subspecies, and varieties. A ‘rare plant’ is
defined as one that is restricted to “a small geographic
area,” or one that occurs “sparsely over a wide area.”
The authors have been cautious to avoid premature
1979
designations of ‘threatened’ or ‘endangered’ in ad-
vance of objective information to defend such a
designation, noting that for some plants “even
information on rarity was scarce.”
Pages 11-49 present an annotated list of 611 plants
arranged in alphabetical order by scientific name.
Where information exists brief notes are supplied for
each plant including “synonyms, references docu-
menting its status as a rare or endangered plant, its
distribution in Canada and Alaska, its distribution in
Ontario, references to published or unpublished
distribution maps and comments on its status in
Canada, special protection and other comments.” The
regimented format of this list suggests a computer
application for processing future lists.
The reader is sure to be alarmed by the length of this
list, which represents approximately 19 percent of the
entire, native vascular flora of Canada. For many of
these plants the center of distribution in Canada, or
the entire Canadian distribution is restricted to
southern Ontario where the pressures for habitat
destruction are severe. For example, in a Canadian
context, many Carolinian plants can be considered
‘endemic’ to Ontario. It may be appealing to suggest
that the massive landscape conversion in southern
Ontario since white settlement is a major reason for
this high incidence of rare plants, but the essential
historical documentation to advance this postulate is
very inadequate.
Pages 50-63, inclusive, contain a bibliography in
excess of 300 references and three appendices.
Appendix | lists 36 plants reported to occur in Ontario
whose status requires verification. Appendix II lists 21
alien or doubtfully native plants. Appendix III
presents the list of rare plants in taxonomic order
following Gray’s Manual of Botany, edition 8.
Fungi: Delight of Curiosity
By Harold J. Brodie. 1978. University of Toronto Press,
Toronto. 131 pp., illus. $10.00.
Here is a book of many charms, but how does one
classify it? It could scarcely be further removed from
the textbook category, although written by a mycol-
Ogist. It is certainly not science fiction, though the
rank beginner or the skeptic might be excused for
thinking so. The author states his aim frankly, to share
his personal feelings of curiosity, excitement and
delight when confronted with the complexity and
perfection of devices in operation among the fungi as
they compete for survival and propagation of their
kind. Written not for students of the fungi, nor even
for those with some background in biology, the book
is addressed to the mycologically uninformed public.
BooK REVIEWS
101
The publication measures approximately 21.7 x
28.1 cm (8 1/2” X 11”) and is neatly bound in soft
cover with a stiff spine. The text is clean, free of typo-
graphical error, and organized into two columns to
facilitate reading. Although the publication includes
both English and French translations, the latter
appears unjustified except for the Introduction on
pages I-11. The bibliography and appendices are
identical duplicates in the two translations while only
very elementary English or French translation skills
are necessary to comprehend either translation of the
annotated rare plant list.
Although traditional views concerning the protec-
tion of endangered species have focused on wildlife in
a quite narrow sense, this publication is evidence of a
broadening concern for the stewardship of our
biological heritage. As the initial statement in a
projected series, The Rare Vascular Plants of Ontario
promises great potential to monitor and to update
periodically a red list of the Canadian flora. In
addition to being an incentive for botanists and
naturalists to continue exploratory work towards a
more comprehensive understanding of endangered
plant life, this report serves as a catalyst to further the
appreciation and the protection of our native flora.
This significant work deserves the serious attention of
all government agencies, private sector organizations
and individuals concerned with biological conserva-
tion and landuse planning in Ontario.
T. J. BEECHEY
Provincial Parks Branch, Ontario Ministry of Natural
Resources, Whitney Block, Queen’s Park, Toronto,
Ontario M7A IW3
It avoids technical jargon, and describes phenomena
of the fungus world in simple terms intelligible to non-
biologists.
Between Preamble and Postcript are eleven un-
related chapters, each complete in itself, describing
strange fungi or curious features of fungus life. The
chapters are bound together very loosely by the fable
of the blind men and the elephant, illustrating the
author’s contention that most of us, limited by
viewpoint and individual powers of perception, fail to
perceive and appreciate many aspects of the natural
world around us.
Some indication of the book’s content may be
gathered from reference to the topic of chapter one:
Gunnery in the Fungus World. The common mold
Pilobolus, a marvel of biological ingenuity, is an
102
example in nature of an automatically controlled
missile. By mysterious manipulations of its chemistry,
and making use of well-known principles of physics,
the fungus has perfected a complex and efficiently
designed apparatus for hurling its projectile (a pellet
of spores) several thousand times its own diameter,
actually directing the aim of the missile with over 80%
accuracy. Equally intriguing is the subject matter of
the other ten chapters.
Not the least of the delights of this book is the
author’s facility with English. With a few strokes of
the pen, he evokes lively mental images of the unseen
fungus world. It is a rare scientist, too, that can align
his thinking with that of ordinary folk, to describe a
fungus as “suggesting a broad vase filled with
bananas,” or again, to depict the fungus-gardening
ant carrying big leaf fragments aloft “like a man
carrying a sheet of plywood.” The author’s first
THE CANADIAN FIELD-NATURALIST
Vol. 93
finding of the Coral Fungus is described in a most
beautiful piece of writing, revealing a fine-tuned
sensitivity to sound, scent, color, and form. Through-
out the book, an unexpected sprinkling of phil-
osophy and folklore, wisdom and humor, make for
enjoyable reading quite apart from the scientific
information imparted.
The 22 black-and-white photographs of fungi have
magnifications ranging from 1/2 x to 10000 x. Also
included are a simple glossary, a selected reference to
books on related topics, and an index.
It is difficult to pigeon-hole this work, but there is
no difficulty in recommending it as a delightful and
unusual addition to one’s natural history library.
SHEILA C. THOMSON
2066 Rideau River Drive, Ottawa, Ontario KIS 1V3
Pollen Flora of Argentina: modern spore and pollen types of Pteridophyta, Gymnospermae,
and Angiospermae
By Vera Markgraf and Hector L. D’Antoni. 1978. Uni-
versity of Arizona Press, Tuscon, Arizona. 208 pp.
US $9.50.
This publication provides photomicrographs, mor-
phologic descriptions, and keys for 374 pollen types. It
divided keys into four plant geographic regions to
facilitate palynologic work in other similar areas. This
text also includes indexes to plant families, plant
species, and common names.
This much needed text will be very useful to people
working in palynology and other disciplines in
botany, not only in South America, but in North
America, Europe, Africa, Asia and Australia.
I. JOHN BASSETT
Biosystematics Research Institute, Canada Agriculture,
Ottawa, Ontario KIA 0C6
Floristics and Environmental Planning in Western New York and Adjacent Ontario —
distribution of legally protected plants and plant sanctuaries
By Richard H. Zander. 1976. Occasional Paper Number 1,
Buffalo Society of National Sciences, Buffalo, New York.
47 pp., illus. Paper US$2.50 + 35¢ handling.
The paper deals with a number of plants that have
been given some protection by New York State law.
All protected species, except for ferns, orchids, and
club mosses are mapped on an individual basis. Many
species are discussed in detail with emphasis on
geographic distribution and preferred habitat.
This publication was obviously written by a
botanist who is concerned about the long-term
protection of rare plants and their habitats. This
reviewer and most readers of The Canadian Field-
Naturalist would agree with this idea.
The paper’s strength lies in its botanical com-
ponents. It is weak inits planning elements. The title is
too broad for the material covered in the small
publication. In reality the paper deals with rare plants
and areas worthy of protection rather than floristics
and environmental planning.
Zander mentions that Section 193.3 of the New
York State Penal Code, “protects certain nature
wildflowers, ferns and shrubs in New York State and
provides for a small fine for picking, transplanting or
removing any of them. However, it affords no
protection to these plants from the owners of the land
upon which they grow.” Later he mentions that “the
recently enacted state Freshwater Wetlands Law...
1979
would give some protection from encroachment to
wetlands over 12.4 acres in extent.” These statements
tantalize. It would have been worthwhile if the
relevant sections of the law had been discussed in
detail. Those who are working towards legislature
changes to protect natural areas, elsewhere, would
have appreciated such information.
The document is liberally illustrated with distribu-
tion maps. Each has a typical dot format which
indicates the presence or absence of a species in a
minor political district (townships, town, or city). It
would have been better if each station of the
uncommon plant had been indicated by a small dot at
the appropriate site location, as was done in Fox and
Soper’s well read papers on Canada’s Carolinean
flora. Unfortunately, several of the maps have no
legend and no reference back to the text page where
some explanation does occur.
Now a few small points. Is Helleborine, Epipactis
helleborine, actually so rare as to bea protected plant
in New York State? In my experience in southern
Ontario it is ubiquitous. In the Regional Munici-
pality of Niagara there are important nature reserves
that do not appear on the list. The Short Hills Nature
Sanctuary, near Forthill, is owned and managed by
the Hamilton Naturalists’ Club. The large, new Short
Hills Provincial Park has extensive blocks of impor-
tant natural areas most of which are protected in the
Master Plan.
Ideally, what would a planner or other decision-
maker need in order to fulfill the expressed desires of
this book? Let’s list a few:
Atlas of Airborne Pollen Grains and Spores in
By Siwert Nilsson, Joseph Praglowski, and Lennart Nilsson.
1977. Ljungforetagen, Orebro, Sweden. 159 pp. US
$37.50: Sw Crs 170.
This is an excellent introductory text on airborne
pollen grains and spores of northern Europe. The
atlas contains descriptions and illustrations of more
than 70 species utilizing light, scanning and trans-
mission electron microscopy. The descriptions are
accompanied by distribution maps and flowering
period for each species. Many of the species described
also occur in Canada and the northern United States.
Freshly collected pollen grains and spores were
generally used. In some cases pollen was sampled
from dried herbarium specimens located in the
Botanical Section of the Swedish Museum of Natural
History. The pollen grains and spores were embedded
in fuschin-stained glycerol jelly and the cover slips on
BOOK REVIEWS 103
(1) A law allowing him to practice environmental
planning in his area of jurisdiction.
(2) The written judgment of reputable botanists on
which plants are worthy of protection and why
they would be protected.
(3) Detailed maps showing the important botanical
areas that need to be protected.
(4) Published data that supports the boundary
delineations of the areas.
(5) Political (public) support when a land-use con-
flict threatens a delineated natural area.
This publication attempts to fulfill several of these
points, most specifically 2 and 3. Each of these efforts
listed above should be in place at a time well before
any on-the-ground activity takes place. For example,
most subdivisions in Ontario usually take 5 years from
when the first approach is made toa municipality until
construction starts. The later an environmentalist
enters the planning flow the less likely is his possibility
of success.
It is the melding of biological data and philos-
ophies into the land-use planning regimes of our
society that 1s now just getting off the ground, or
should I say on the ground. Publications such as this
one are efforts in that direction.
PAUL F. J. EAGLES
Faculty of Environmental Studies, University of Waterloo,
Waterloo, Ontario N2L 3GI
Northern Europe
the slides sealed with paraffin wax. Preparations for
scanning, transmission electron microscopy are clear-
ly explained in the text.
It is unfortunate that there was no description of the
equipment used to collect airborne pollen and spores.
The amounts of airborne pollen and spores can be
summarized more specifically by using one of the
collecting devices now used in several countries. |
doubt, for example, that the very minute airborne
pollen grains from dandelion, Taraxacum spp. or
from goldenrod, Solidago spp., would be caught in
very large amounts.
I. JOHN BASSETT
Biosystematics Research Institute, Canada Agriculture,
Ottawa, Ontario KIA 0C6
104
Orchid Biology: reviews and perspectives, I
Edited by Joseph Arditti. 1977. Cornell University Press,
Ithaca. 310 pp. US $29.50.
Biology is a wide field of study, even if all the
organisms studied belong to a single plant family.
Even so, its meaning has to be stretched to cover the
range of topics in this book. One third of the text is
devoted to “A manual for clonal propagation of
orchids by tissue culture.” In this chapter, J. Arditti
has gathered together all the techniques brought to his
attention by the cut-off date: December 1974. Not
only does he cite the original papers but also provides
comments based on his own experience. As an
annotated review of the literature it should be of some
value to people in the area. The other articles seem of
less value, either because they cover little new ground
or because there is so little good information
available. The account of the fossil history of the
Orchidaceae appears to fall in the first category. To
people like myself who have not read the Schmids’
previous articles on the topic, published in 1973 and
1974 and cited here, their article is interesting and
provides a useful warning on the perils of using
ENVIRONMENT
Nature Quizzes for Canadians
By V. and B. McMillan. 1976. Douglas, Vancouver. 144
pp., illus. Paper $3.95.
The impetus behind this book evolved from a desire
to assist Canadians in learning about their country’s
natural environment. I must admit, there are some
interesting and informative chapters; however, the
text is poorly organized and filled with errors.
The numbering systems employed to differentiate
the questions are very poor. In many cases the
numbers used to label the diagrams are also designed
to double as question numbers. On page 89 the first
seven questions are labelled alphabetically froma to g,
while the remaining questions in the chapter read as
numerics (8--20). Very confusing.
In the chapter examining the reader’s knowledge of
‘Immature Insects,’ pictures of the adult forms, which
the reader is asked to match up with the immature
stages, are completely missing. On page 90 (question
11) the authors make reference to a diagram that does
not exist.
I also found a question (#5, page 15) where the
THE CANADIAN FIELD-NATURALIST
Vol. 93
secondhand sources of information. I am doubtful,
however, of its value to students of orchids. The article
on anthocyanins of the Orchidaceae falls into the
second class: not enough information is available to
justify the length of this section. Two other articles fall
outside the bounds of orchid biology to my mind: an
account of his involvement with orchids by R. E.
Holttum and a discussion of Rumphius’ contribution
to orchidology by H.C. D. de Wit. I enjoyed them
both as general reading, although that on Rumphius
is, in my judgment, rather long. This article and the
others would benefit from a more severe editorial
hand. As it stands, the section on tissue culture
techniques should earn this book a place on the
shelves of orchid breeders and the title will recom-
mend the book to ardent orchid devotees. It is perhaps
only fair to admit that I myself am not one.
M. BARKWORTH
Biosystematics Research Institute, Ottawa, Ontario
KIA 0C6
wrong answer, followed by a correct explanation, was
recorded, thus causing a contradiction, which to a
novice might prove confusing. On page 41 (question
20) the reader is asked to give the name of a northern
freshwater fish belonging to the Gadidae. Directly
below the question, not in the answer section where it
belongs, is the answer, a labelled diagram of a burbot.
There is no consistency in the usage of nomen-
clature. I found three incorrect spellings: the red
admiral butterfly (Vanessa atlanta) is not Vanessa
atalenta (page 38); the moose (A/ces alces) is not Alcea
alcea (page 40); the muskrat (Ondatra zibethicus) is
not Ondatra zibethica (page 49).
I feel that the concept is sound, and that the text has
possibilities, but until the book is properly organized
and the errors in composition are corrected, the book
is not worth buying.
PAUL A. GRAY
Ontario Ministry of Natural Resources, Box 89, Cochenour,
Ontario POV ILO
1979
Book REVIEWS 105
The Shetland Way of Oil. Reactions of a small community to big business
Edited by John Button. 1976. Thuleprint Ltd., Sandwick,
Shetland. 134 pp. £2.40.
This is an admirable little book. Although it deals
with events in a faraway place, under a jurisdiction
and an administrative system that differ from those in
Canada, it offers much sensible advice, as well as
stimulating examples of success, to all who care for
more in the world they live in than just their own
financial prosperity.
The most powerful reason for the recent setbacks
suffered by the “environmental movement” has been
the sustained demand for oil, despite the massive price
increases imposed by the OPEC countries. Part of the
western reaction was to intensify the search for other
supplies. Their most substantial early success was the
discovery of large quantities of oil and gas in the
northern North Sea and the development of tech-
nology to extract oil from under the seabed at
unprecedented depths. The nearest deep-water harbor
to some of the most promising oil fields was found at
Sullom Voe, off the Mainland of Shetland. So there
the oil industry descended, with plans for the
development of a storage facility for crude oil brought
ashore by pipeline and later establishment of a
refining plant too.
The fourteen chapters of this book, each by a
different author, half of them local men, deal chiefly
with the responses of the local communities to the
potent and unexpected combination of opportunities
and hazards. By great good fortune, the expertise and
the affluence of the oil companies and their supporters
in distant arenas of government did not altogether
overwhelm the local talent. The Zetland County
Council (now the Shetland Islands Council) quickly
devised an Interim Development Plan. It succeeded in
obtaining legislation (the Zetland County Council
Act of 1974) which gave it powers exceptional for a
local authority. The council now acts as the port and
harbor authority for Sullom Voe; can acquire land for
oil-related development within the designated area;
issues licences to dredge (and may refuse to do so) and
licences construction work within the three-mile
territorial limit; may take shares in commercial
undertakings; and may establish a Reserve Fund.
With so strong a jurisdictional base the local people
have been well placed to exercise control over
undesirable development and have succeeded in doing
so to a remarkable extent.
For naturalists the chapters by Peter Kinnear on
“Birds and Oil” and by Brian John on “The Milford
Haven Experience” are of greatest interest. Kinnear
writes “The value that is placed on birds and other
wildlife, which do not provide some immediate
financial return, is to a large extent subjective. For
many people the value might seem sufficient, but it
cuts little ice with either Government or developers,
who require a more qualified scientific appraisal. . . .
One way of assessing the importance of Shetland’s
birdlife is to consider the number of different species
and to see what proportion of the total British stocks
they constitute.” Fortunately for his chosen approach,
those numbers are known and for several species do
make up an impressive fraction of the British stock:
90% of the Great Skuas (Stercorarius skua) (or 50% of
all those in the northern hemisphere) and 70% of
Arctic Skuas (S. parasiticus); 90% of the Whimbrel
(Numenius phaeopus), 1/2 to 2/3 of the Red-necked
Phalaropes (Phalaropus lobatus), 1/3 to 1/2 of the
Red-throated Divers (Gavia stellata) and so on, with
nationally significant stocks of at least 24 species.
Most of those species are marine and so directly at risk
from oil pollution, from pipeline leaks, spills on shore,
bilge discharge and tank washing by tankers and,
eventually no doubt, tanker collisions or groundings.
To look at the probability Kinnear turns to the
detailed experience gained over the last fifteen years
or so in and around Milford Haven, in west Wales, a
major reception area for large tankers and “widely
regarded as the cleanest oil port in the world.” Milford
Haven handled 53.1 million tons of oilin 1973. Sullom
Voe is expected to handle 50 to 150 million tons
annually over the next twenty years or so. In the
eleven years 1963-1973 Milford experienced an
average of 53 spills a year. Sullom can thus expect a
similar or larger number, amounting to between 400
and 1200 tons a year, excluding any major accident.
But, whereas Milford Haven is subjected to massive
tidal flushing, Sullom Voe has only one eighth of
Milford Haven’s tidal velocity and only a fraction of
its water is changed each tidal cycle. Building up of oil
and other pollutants seems inevitable; and a major
tanker accident is likely too, because manoeuvring -
very large tankers through Yell Sound “will always be
a hazardous affair, particularly during fog and severe
weather.” Kinnear, however, sees the greatest danger
to birds as likely to result from the continuing practice
of discharging oily ballast water at sea (in order to
save time at the terminal), an activity which it is often
hard to detect and prove. He complains that British
regulations governing pollution control are in dire
need of change. “Although various laws restrict the
discharge and dumping of oil in the sea, once there it is
incumbent on no-one to do anything about it.”
What lessons can naturalists in Canada learn from
this book? First, in many areas now being explored for
oil we still lack the basic inventories to establish
numerically what birds are at risk, although great
progress has been made along the eastern Arctic and
106
Atlantic coasts and offshore in the last decade. That
task must be completed and the results made known
to governments, the oil industry and the public,
whether or not they want to hear about them. Second,
the best hope for effective actions on behalf of wildlife
lies in power being given to local people and in their
vigilant and responsible use of those powers. (The
Inuit seem to understand this more clearly than many
people elsewhere.) Third, wildlife interests are most
unlikely to be sufficient to prevent major economic
developments, the equivalent of North Sea oil, from
taking place; but they can and should make an
important contribution to keeping the activities of
exploiters cleaner than the latter would like, though
not as clean as the birds require. So longas oil is taken
THE CANADIAN FIELD-NATURALIST
Vol. 93
from under the sea or carried across it, birds will be the
losers.
“So we return to the moral question and the
personal value each one of us places on birds and
wildlife. While you decide how you are going to play
your part in changing things, spare a thought for the
vast numbers of birds which migrate at night and each
year die in thousands, attracted by and consumed in
the giant flames burning from the rigs across the
North Sea” (page 99).
HuGH BoyD
Canadian Wildlife Service, Environment Canada, Ottawa,
Ontario KIA 0E7
EZAIM: écologie de la zone de l’aéroport international de Montréal — Le Cadre d’une recherche
écologique interdisciplinaire
Par P. Dansereau. 1976. Les Presses de l’Université de
Montréal, Montréal. Polycopié. 343 pp.
Comme a déja dit Pun des chefs d’€quipe impliqueé
dans cette étude écologique de la zone de l’a€roport
devenu Mirabel, étude a priori multidisciplinaire: “on
a limpression qu'il n’y a que Dansereau qui fait du
multidisciplinaire.” Ce commentaire souleéve a lui seul
Yampleur du probleme de la communication entre des
spécialistes de disciplines aussi variées que la psy-
chologie, la geomorphologie, la zoologie. Ce méme
probleme cede cependant le pas a la science elle-méme
a lintérieur d'une méme spécialiteé.
Ayant participé a cette étude, j'ai pu me rendre
compte que le travail d’équipe fonctionnait au niveau
de chaque équipe dans leur discipline propre, mais que
le travail d’intégration multidisciplinaire ne semblait
quétre le fruit d'une démarche personnelle, celle de
Dansereau. Ce dernier se veut cependant plus positif
quant a la possibilité et la réalisation d’un projet
multidisciplinaire et complexe a souhait.
L’étude EZAIM a été entreprise apres le début des
travaux de construction et d’ameénagement de l’aéro-
port, ce qui souléve peut-é€tre son caractere secondaire
aux yeux des intéréts €conomico-politiques, probleme
soulevé par Dansereau lui-méme en fin d’ouvrage
d’ailleurs.
Le plan de recherche envisagé a été calqué sur
’écosysteme lui-méme, a savoir les différents niveaux
trophiques entre lesquels ont lieu des interactions. Les
€quipes étaient donc divisées en géomorphologie,
écologie végétale, écologie animale (mammiferes,
olseaux), géographie humaine (utilisation des terres,
industrie et urbanisation), psychiatrie sociale, sans
compter les échelons nécessaires de coordination et
d’administration.
Dansereau insiste sur le fait que cette recherche se
veut une expérience, multidisciplinaire soit, mais
expérience intégrée par l’auteur dans le contexte d’une
“philosophie écologique” quil prone. Le chapitre 2 est
peut-étre a ce point de vue le plus important, car
Yauteur y expose les définitions de sa démarche qui
doit assurer la cohérence a la recherche (ou a toute
recherche) qui s’attaque finalement a des €cosystemes
tres variés.
La définition élargie qu'il donne a l’écosystéme est
la suivante: “Un espace limité ot le cyclage des
ressources a travers un ou plusieurs niveaux tro-
phiques est effectué par des agents plus ou moins fixés
et nombreux, utilisant simultanément et successive-
ment des processus mutuellement compatibles qui
engendrent des produits utilisables 4 courte ou longue
échéance.”
Se basant sur cette hypothése de travail, et par une
description des relations qui interviennent entre les six
niveaux trophiques (minéro-, phyto-, zootrophie
(herbivorie, carnivorie), investissement et controle)
dans des écosystemes types, c’est-a-dire par une
analyse écosystématique des paysages, il parvient a
définir tout paysage d’un point de vue écologique.
Cette analyse consiste en un inventaire des ressources
présentes a chaque niveau trophique et des agents et
processus ou forces motrices ayant court dans la
transformation d’un niveau a l'autre ou d’un paysage a
l'autre. Elle implique également la compréhension de
l'aspect dynamique du cyclage dans chaque paysage
pour mieux situer la stratégie régionale d’un point de
1979
vue écologique, et mener ainsi a la construction de
matrices d’impact qui devraient guider la planifi-
cation.
Mais non content de l’analyse €cosystématique de la
zone de Mirabel, Dansereau débouche naturellement
sur Vuniversel, soit lorsquwil présente les régions
phytotrophiques, dont celles applicables a Mirabel,
soit lorsquil présente son nouveau systeme de
classification des terres basé sur l’escalade de impact
humain. Et ce sont justement ces visées universalistes
qui font de cet ouvrage de Dansereau un outil de plus
pour les études a caracteére multidisciplinaire.
Toute cette philosophie écologique peut se résumer
graphiquement par un “boule-de-fléches” (sphére qui
contient les circuits de ressources, agents, processus et
produits) qui illustre sa démarche. Cette démarche n’a
pas été comprise par tous les participants du projet
EZAIM, ou selon lexpression de Dansereau “a
souvent été perdue de vue.”
Du point de vue strictement de la présentation, les
erreurs typographiques sont rares. Certaines figures
ou tableaux sont incomplets: la légende de la figure 23
nest pas rapportée enti¢rement au tableau XIII tel
que mentionneé. Le style est simple et clair, sice n’est le
vocabulaire scientifique propre au domaine du
botaniste qui transpire tout au long de l’ouvrage.
BooK REVIEWS 107
On y trouvera aussi une autocritique, sirement pas
assez vive a mes yeux et a ceux de certains autres
participants, du projet EZAIM tel qu'il avait été
con¢u par l’auteur du présent ouvrage et tel qu’ila été
réalisé. Globalement, Vouvrage rend davantage
compte de ce que devrait étre une recherche €colo-
gique interdisciplinaire a partie d'un modéle, plutdt
que d’étre une synthese des résultats écologiques
recueillis par chaque équipe. Ce qui retiendra notre
attention sera ce modéle €cosystématique de Dan-
sereau, sa philosophie qui permettrait d’analyser tous
les types de paysage selon un point de vue €cologique,
y compris les paysages humains et il faut, je crois
rendre hommage a Dansereau pour sa vision syn-
thétique du cadre qu'il propose.
On ne saurait trop recommander la lecture de cet
ouvrage aux aménagistes et aux spécialistes a tous les
niveaux trophiques, tant naturels qu’artificiels ou
humains, tout au moins pour les changements dans les
valeurs sociales qu'il préne pour les développements
d’ordre socio-économiques a venir.
ANDRE CYR
Département de Biologie, Université de Sherbrooke, Sher-
brooke, Québec JIK 2RI
Outdoors Canada. A unique and practical guide to our wilderness and wildlife
Edited by Douglas R. Long. 1977. Reader’s Digest Canada
Ltd. and the Canadian Automobile Association, Mon-
treal. 383 pp., illus. $21.77.
Outdoors Canada is intended to assist Canadians to
enjoy their out-of-doors. The book not only tells of
the out-of-doors (mammals, birds, plants), but it also
explains how to cope in the out-of-doors (camping,
canoeing, backpacking). Photographs demonstrate
what can be seen, while numerous drawings illustrate
and explain the book’s “how to” information.
Not all of the information in the section on wildlife
is entirely accurate. Some notable inaccuracies would
include these statements: “many fur bearing animals
are near extinction” (p. 15); “most great whales are
endangered species” (p. 13); “the distribution of
porcupine is south of the treeline” (should read mainly
within the predominantly hardwood forest areas).
The section is far from complete but does present an
interesting capsulized summary of most groups of
animals found in Canada.
As noted within the text, birds are animals and are
included within the animal section. The treatment
given to birds is less complete than that given to
mammals. Again some of the book’s statements are
questionable. One such statement to which some
persons might take exception would be, “the whoop-
ing crane is North America’s rarest bird.” The section.
on birds will not serve as a field guide; however, it is
enjoyable reading.
Whereas for the section on Animals the photo-
graphs tend to supplement the text, the reverse
treatment is given to the section on Plants: photo-
graphs are supplemented by the text. Also, the Plant
section tends predominantly to focus on flowering
plants and generally those that are more common.
Again this section will not replace a field-guide
treatment as is popularly used by naturalists. Unlike
the Animal section, this section appears disorganized.
The section on Camping covers the whole gambit of
outdoor travel. This section is generally well or-
ganized, well presented, and contains much good .
advice. One comment I would caution the reader
about is this: “to keep your feet dry in leaky boots orin
shoes that are not waterproof, wear plastic bags over
108
socks.” If the weather is cold, perspiration within the
plastic bags could result in very cold feet (possibly
frozen). As the book points out, “experience is
probably the surest guide.”
As surely as rules are made, there are exceptions.
The section, “Outdoors: Just for Fun,” suggests it is a
good idea to stay with your canoe after it is capsized.
But common sense should prevail and when swamped
or capsized in white water, sometimes it is best to leave
the canoe. On occasion, people have been known to
get pinned between their canoe and rocks, resulting in
serious injuries.
The last 49 pages of Outdoors Canadais a series of
regional (Canadian) maps. These pinpoint the loca-
tions of various outdoor areas in Canada. A brief note
THE CANADIAN FIELD-NATURALIST
Vol. 93
explains the features of specific sites. This section will
be of assistance to those persons planning a Canadian
out-of-doors vacation.
Generally speaking, Outdoors Canada is very much
a Reader’s Digest-type book. It is a brief summary of
material rather than a complete treatment. The book
covers the whole spectrum of the outdoors and
admirably does not get bogged down in any one
section. For the inexperienced outdoors-person, it 1s
an excellent buy.
PETER CROSKERY
Ontario Ministry of Natural Resources, Ignace, Ontario
POT ITO
Why Big Fierce Animals Are Rare. An ecologist’s perspective
By P. Colinvaux. 1978. Princeton University Press, Prince-
ton, New Jersey. viii + 256 pp. $9.50.
In an informative and logical fashion Colinvaux
presents the rudiments of ecology, and endeavors
...“to trace the status of the ecologist’s quest” ina
thorough examination of the numerous riddles of
nature. Without obscuring the text with highly
technical terms that might discourage the novice, he
reveals concerns of modern ecologists, concerns
nurtured by pioneers like Charles Darwin, G. F.
Gause, and Charles Elton.
In 18 chapters he outlines the niche, biogeography,
aquatic-terrestrial-atmospheric cycles, succession, co-
existence, behavior, species diversity, environmental
stability, and the evolution of man’s role in nature.
Page by page the ecological story unfolds as the
author describes the efforts of men like Alphonse de
Candolle, the last man singly to attempt to describe all
of the known plants in the world; Vladimir Koppen,
the climatologist who mapped world weather using
vegetation zones; C. Hart Merriam, the explorer who
devised the life zone concept; and Tansley, the English
botanist who coined the word ‘ecosystem.’
The subject matter is uniquely accented with a
number of interesting anecdotes. For example, in
chapter 3, entitled “Why Big Fierce Animals Are
Rare,” Colinvaux exposes the myth of 7yrannosau-
urus rex amidst a discussion of food-chain dy-
namics. He also dispels the fantasies of hope that
evolved from the green revolution in a few short
sentences: “It is a myth that is probably as impossible
to eradicate as the myth that Tyrannosaurus rex was
a ferociously active predator. But myth it is. Algae are
not more productive than other plants.”
I was particularly impressed with the final chapter,
“The People’s Place,” in which the author recon-
structs the story of man’s climb to environmental
dominance. But more than that, Colinvaux stresses
the importance of man’s dynamic role in the complex
ecosystems of the earth, a role that beginning
ecologists must understand in order to develop an
appreciation for the relations among other living
organisms and their environment.
The text is greatly enhanced by a section on
ecological reading, which lists many excellent papers
on the numerous subjects of the science. A useful
index is also included. Unfortunately, the title of the
book, Why Big Fierce Animals Are Rare, is mis-
leading; in my opinion the title suggests that the book
is about predator ecology, not a general overview of
ecology. Nonetheless, it is well written, and is as
complete a book as one would expect at the
introductory level.
PAUL A. GRAY
Ministry of Natural Resources, Box 89, Cochenour, Ontario
POV ILO
1979
OTHER
Northern Vagabond. The life and career of J. B.
By Alex Inglis. 1978. McClelland and Stewart, Toronto.
256 pp., illus. $14.95.
The author begins this interesting book by des-
cribing Joseph Burr Tyrrell’s famous 1893 3000-mile
trip through the central Barren Lands of Canada —
one of the last great feats of exploration in Canada’s
history. When Tyrrell’s party of eight set out by canoe
from Fort Chippewyan they were not certain whether
the Dubawnt and Thelon rivers, along which they
planned to paddle, would carry them to Hudson Bay
or to the Arctic Ocean; if the latter, their chances of
returning alive were slim. Nor were they sure they
would encounter game, largely Caribou, to feed them.
Their inland exploration ended successfully at
Chesterfield Inlet on Hudson Bay, but their trip was
by no means over. They had yet to make their way
through blizzards and ice along the coast for 400 miles
to Fort Churchill, and from there walk and travel by
dog sled the 900 miles to Winnipeg and the railway.
Tyrrell undertook this voyage while he was a
geologist with the Geological Survey of Canada, a
position he held from 1881 to 1898 although he had
little formal training as a scientist. He spent his
summers exploring the wilds of western and northern
Canada, his winters in Ottawa writing reports on his
geological and other findings. He enjoyed his field
work especially (in one week in Alberta he discovered
the largest deposit of prehistoric bones and the
greatest deposit of workable coal to be found in
Canada). He is probably best known scientifically for
his postulation, based on rock striations, that during
NEW TITLES
Zoology
Animal facts and feats. 1977. By Gerald L. Wood. Revised
edition. Sterling, New York. 256 pp., illus. US $14.95.
Animals in peril. A guide to the endangered species of
Canada and the United States. 1978. By David C. Grainger.
Macmillan, Toronto. 192 pp., illus. $14.95
The beetles of North America. 1977. By Richard Head-
strom. Barnes, Cranbury, New Jersey. 448 pp., illus. US
$17.50.
The California quail. 1978. By A. S. Leopold. University of
California Press, Berkeley. xx + 281 pp. US $14.95.
Book REVIEWS
109
Tyrrell
the Ice Age Canada was not covered by a single
massive ice field which advanced and retired once ona
long front, but by several glaciers which originated
from separate centers and had advanced and retreated
on several occasions. During his lifetime, besides his
geological reports, he also wrote dozens of books and
articles on such topics as Samuel Hearne, David
Thompson, Canadian history, and mining in Canada.
In 1899 Tyrrell left the Survey because he had been
denied promotion, and moved to the Klondike as a
Mining Consultant. He lived there in the midst of the
gold rush until 1905, learning so much about gold that
the rest of his career was centered on this mineral, in
part as a goldmine manager near Kirkland Lake,
Ontario. When he retired a wealthy man, he turned to
managing an apple orchard on the Rouge River,
where many of us who lived in Toronto at the time
were privileged to meet him. He died in 1957 at the age
of 99.
This book isa timely one, dealing as it does with one
of our first native-born natural scientists. It not only
follows Tyrrell’s life in detail, but it gives us a glimpse
of the milieu in which an early government scientist
worked (The Ottawa Field-Naturalists’ Club meet-
ings for example offered Tyrrell and his friends a
welcome diversion from their work). We are indebted
to Inglis for bringing the life of this remarkable
Canadian to our attention.
ANNE INNIS DAGG
Box 747, Waterloo, Ontario N2J 4C2
A changing world for wildlife. 1977. By G. S. Fichter.
Golden/ Western, New York. 141 pp., illus. US $8.95.
Coyotes: biology, behavior and management. 1978. Edited
by M. Bekoff. Academic, New York. 400 pp. US $34.50.
+Endangered birds. Management techniques for preserving
threatened species. 1978. Edited by Stanley A. Temple.
University of Wisconsin Press, Madison. xxiii + 466 pp.,
illus. US $9.50.
+L’entomologiste amateur. 1977. Par A. Villiers. Savoir en
histoire naturelle xxvii. Lechevalier, Paris. 248 pp. illus. 90F.
Great Canadian animal stories. 1978. Edited by Muriel
110
Whitaker. Hurtig, Edmonton. 232 pp., illus. $12.95.
*Mammals of the palaearctic region: a taxonomic review.
1978. By G. B. Corbet. British Museum (Natural History).
Cornell University Press, Ithaca. 314 pp., illus. US $38.50.
*Nesting ecology of Canada geese in the Hudson Bay
Lowlands of Ontario: evolution and population regulation.
1977. By Dennis G. Raveling and Harry G. Lumsden. Fish
and Wildlife Report No. 98. Ontario Government Book-
store, Toronto. v + 77 pp. Paper $3.50.
The response of peregrine falcons (Falco peregrinus) to
aircraft and human disturbance. 1977. By Jim Windsor.
Canadian Wildlife Service, Ottawa. 87 pp.
The Running Press book of turtles. 1977. By Richard E.
Nicholls. Running Press, Philadelphia. 150 pp., illus. Paper
US $4.95.
The view from the oak: the private worlds of other
creatures. 1977. By Judith and Herbert Kohl. Sierra Club/
Scribners, San Francisco. 112 pp., illus. Cloth US $8.95;
paper US $4.95.
Water birds of California. 1977. By H. L. Cogswell and G.
Christman. University of California Press, Berkeley. 399 pp.
Paper US $5.75.
Wild birds of the Americas. 1977. By Terence Michael
Shortt. Houghton Mifflin, Boston. 272 pp., illus. US $14.95.
Botany
+Ferns of the Ottawa district. 1978. By William J. Cody.
Agriculture Canada Publication 974. Supply and Services
Canada, Hull. 112 pp., illus. $3.25 in Canada; $3.90
elsewhere.
A field guide to edible wild plants of eastern and central
North America. 1978. By L. Peterson. Houghton Mifflin,
Boston. 384 pp. US $8.95.
Grass: the everything, everywhere plant. 1977. By Augusta
Goldin. Nelson, New York. 176 pp., illus. US $7.95.
Mushrooms of North America. 1977. By O. K. Miller, Jr.
Chanticleer/Dutton, New York. 368 pp., illus. Paper US
$8.95.
+Native trees and shrubs of Newfoundland and Labrador.
1978. By A. Glenn Ryan. Newfoundland Department of
Tourism, St. John’s. 117 pp., illus. Free.
tNorth American forest lands at latitudes north of 60 degrees.
1978. By various authors. Proceedings of a symposium,
University of Alaska, 19-22 September, 1977. School of
Agriculture and Land Resources Management, University of
Alaska, Fairbanks. 332 pp., illus. Free.
Syrup trees. 1978. By Bruce Thompson. Walnut Press,
Fountain Hills, Arizona. US $6.95.
Wild food plants of Indiana and adjacent states. 1977. By
Alan and Sue McPherson. Indiana University Press,
Bloomington. viii + 215 pp., illus. Paper US $4.95; cloth US
$12.50.
Environment
Biological data in water pollution assessment: quantitative
and statistical analyses. 1978. Edited by K. L. Dickson,
THE CANADIAN FIELD-NATURALIST
Vol. 93
John Cairns, Jr., and R. J. Livingston. Special Technical
Publication 652. American Society for Testing and
Materials, Philadelphia. 193 pp. US $17.50 (less 20% to
ASTM members).
Biological nomenclature. 1978. By C. Jeffrey. 2nd edition.
Crame, Russak, New York. viii + 72 pp. US $11.50.
The breakdown and restoration of ecosystems. 1978. Edited
by M. W. Holdgate and M. J. Woodham. NATO Conference
Series I: Ecology, Volume 3. Plenum, New York. 506 pp. US
$36.
Concepts of applied ecology. 1978. By R.S. DeSanto.
Springer-Verlag, New York. 320 pp. Paper US $9.80.
+Environmental role of nitrogen-fixing blue-green algae and
asymbiotic bacteria. 1978. Edited by U. Granhall. Ecologi-
cal Bulletins NFR 26. Swedish National Science Research
Council, Stockholm. 400 pp., illus. 100SwCr.
Estimating the hazard of chemical substances to aquatic life.
1978. Edited by John Cairns, Jr., K. L. Dickson, and A. W.
Maki. American Society for Testing and Materials, Philadel-
phia. 283 pp. US $19.50 (less 20% to ASTM members).
Evolutionary ecology. 1978. By E. R. Pianka. Harper and
Row, New York. 384 pp. US $16.95.
The green world: an introduction to plants and people.
1978. By Richard M. Klein. Harper and Row, New York.
512 pp. US $14.95.
The management of estuarine resources in Canada. 1978. By
Irving K. Fox and J. P. Nowlan. Report No. 6, Canadian
Environmental Advisory Council, Ottawa. 51 pp. Free.
Mexican wilderness and wildlife. 1978. By Ben Tinker.
University of Texas Press, Austin. xii + 132 pp., illus. US
$9.95.
Reports of the first and second meetings of publie interest
groups with the Canadian Environmenta! Advisory Coun-
cil. 1978. By anonymous. Report No. 7, Canadian En-
vironmental Advisory Council, Ottawa. 124 pp. Free.
Resources, environment and economics. 1978. By R. U.
Ayres. Wiley-Interscience, Somerset, New Jersey. 240 pp.
US $24.95.
Sourcebook on the environment. A guide to the literature.
1978. Edited by K. A. Hammond, G. Macinko, and W. B.
Fairchild. University of Chicago Press, Chicago. x + 614 pp.
US $22.
Water and society: conflicts in development. Part 1. The
social and ecological effects of water development in
developing countries. 1978. Edited by Carl Widstrand.
Pergamon, New York. 134 pp., illus. US $20.
Miscellaneous
Carbon dioxide, climate and society. 1978. Edited by Jill
Williams. Proceedings of an IIASA Workshop, 21-24
February, 1978. Pergamon, New York. 304 pp. US $30.
Earth observation systems for resource management and
environmental control. 1978. Edited by Donald J. Clough
and L. W. Morley. Plenum, New York. 475 pp. US $42.50.
72)
Journey through the universe. An introduction to as-
tronomy. 1978. By T.L. Swihart. Houghton Mifflin,
Boston. xvi + 366 pp., illus. US $15.95.
The mountains of Canada. 1978. By Randy Morse. Hurtig,
Edmonton. 144 pp., illus. $29.95.
Quaternary geology: a stratigraphic framework for multi-
disciplinary work. 1978. By D. Q. Bowen. Pergamon, New
York. 224 pp., illus. Cloth US $30; paper US $12.50.
Manual on water. 1978. Edited by C. E. Hamilton. Special
Technical Publication 442A. American Society for Testing
and Materials, Philadephia. 471 pp. US $28.50 (less 20% to
ASTM members).
BoOK REVIEWS 111
Correction
The listings of “Zoology New Titles,” Canadian Field-
Naturalist 92(3): 316, 1978, should read as follows:
Biology of bats, volume 3. 1977. Edited by W. A. Wimsatt.
Academic, New York. 651 pp., illus. US $59.
Biology of bryozoans. 1977. Edited by R. M. Woolacott and
R. L. Zimmer. Academic, New York. 556 pp., illus. US $35.
tavailable for review
*assigned for review
Instructions to Contributors
Content
The Canadian Field-Naturalist is a medium for the
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naturalists or field-biologists reporting observations and
results of investigations in any field of natural history
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(1978) available from the American Institute of Biological
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TABLE OF CONTENTS (concluded)
News and Comments 87
Book Reviews
Zoology: A guide to bird finding east of the Mississippi — The birds of New Brunswick — Mountain 91
monarchs: wild sheep and goats of the Himalaya — Précis de zoologie: vertébrés. 1: reproduc-
tion, biologie, évolution et systématique. Oiseaux et mammiferes — Ways of the six-footed —
Birds of southeastern Michigan and southwestern Ontario — Watching sea birds — Bird hazards
to aircraft: Problem and prevention of bird/ aircraft collision — Bears: their biology and man-
agement — The earthworms (Lumbricidae and Sparanophilidae) of Ontario — The Passenger
Pigeon: its natural history and extinction — The bird finder’s 3-year note book.
Botany: The rare vascular plants of Ontario/Les plantes vasculaires rares de Ontario — Fungi: 100
delight of curiosity — Pollen flora of Argentina: modern spore and pollen types of Pteridophyta,
Gymnospermae, and Angiospermae — Floristics and environmental planning in western New
York and adjacent Ontario: distribution of legally protected plants and plant sanctuaries —
Atlas of airborne pollen grains and spores in northern Europe — Orchid biology: reviews and
perspectives, I.
Environment: Nature quizzes for Canadians — The Shetland way of oil: reactions of a small com- 104
munity to big business — EZAIM: écologie de la zone de l’aéroport international de Montréal:
le cadre d’une recherche écologique interdisciplinaire — Outdoors Canada: a unique and prac-
tical guide to our wilderness and wildlife — Why big fierce animals are rare: an ecologist’s
perspective.
Other: Northern vagabond: the life and career of J. B. Tyrrell 109
New Titles 109
Mailing date of previous issue 8 January 1979
1978 Council — The Ottawa Field-Naturalists’ Club
President: R. A. Foxall E. Beaubien C. Gruchy
Vice-President: R. Taylor C. Beddoe P. Hall
Treasurer: B. Henson W. J. Cody V. Hume :
Recording Secretary: D. R. Laubitz Hones BS aaa
Corresponding Secretary: A. Armstrong A. Dugal J. K. Strang
C. Gilliatt EVE) Dy iiodd
Those wishing to communicate with the Club should address correspondence to: The Ottawa-Field Naturalists’ Club,
Box 3264, Postal Station C, Ottawa, Canada K1Y 4J5. For information on Club activities telephone (613) 722-3050.
THE CANADIAN FIELD-NATURALIST Volume 93, Number 1 1979
President’s Message
One hundred years in perspective — the changing roles and objectives of
The Ottawa Field-Naturalists’ Club ROGER A. FOXALL
Guest Editorial
Biology — the unknown science? YORKE EDWARDS 6
Editorial
The Canadian Field- Naturalist — the status quo or a new direction?
LORRAINE C. SMITH 10
Articles
Demographic and dietary responses of Red-tailed Hawks during a
Snowshoe Hare fluctuation
ROBERT S. ADAMCIK, ARLEN W. TODD, and LLOYD B. KEITH 16
The Oriskany sandstone outcrop and associated natural features,
a unique occurrence in Canada
DIANNE FAHSELT, PAUL MAYCOCK, GORDON WINDER and CRAIG CAMPBELL 28
Breeding areas and overnight roosting locations in the northern range of the Monarch
Butterfly (Danaus plexippus plexippus) with a summary of associated migratory
routes FRED A. URQUHART and NORAH R. URQUHART 41
Reproductive biology of the Big Brown Bat (Eptesicus fuscus) in Alberta
DAVID B. SCHOWALTER and JOHN R. GUNSON 48
Wild Mallard stocking in a large marsh habitat ROBERT O. BAILEY 5)
Notes
Recent collections of the Black Redhorse, Moxostoma duquesnei, from Ontario
EDWARD KOTT, ROBERT E. JENKINS, and GREGORY HUMPHREYS 63
Xanthochroism in the Evening Grosbeak CHRISTOPHER W. HELLEINER 66
Interaction between a Long-tailed Weasel and a Snowy Owl PETER) @ Boxe 67
Black-necked Stilts nesting near Edmonton, Alberta ;
DICK DEKKER, ROBERT LISTER, TERRY W. THORMIN, D. V. WESELOH, and LINDA M. WESELOH 68
Common Garter Snake predation on Robin nestlings KATHY MARTIN __70
Distribution and movements of Selkirk Caribou, 1972-1974 DAVID J. FREDDY 71
Effects of fire on the location of a Sharp-tailed Grouse arena
DONALD A. SEXTON and MURRAY M. GILLESPIE 74
Brewer’s Blackbird breeding in the Northwest Territories PHILIP H. R. STEPNEY 76
A trap to measure Populus and Salix seedfall JOHN C. ZASADA and ROSEANN DENSMORE 1
Examination of overwintering adult carabid beetles for associated mites
J. E. OLYNYK and R. FREITAG 79
Ring counts in Salix arctica from northern Ellesmere Island D. B. O. SAVILE 81
Giant Cow Parsnip (Heracleum mantegazzianum) on Vancouver Island, British Columbia
NEIL K. DAWE and ERIC R. WHITE 82
Nesting of Horned Puffins in British Columbia
R. WAYNE CAMPBELL, HARRY R. CARTER, and SPENCER G. SEALY 84
concluded on inside back cover
ISSN 0008-3550
The CANADIAN
FIELD-NATURALIST
Published b
y THE OTTA
WA FIELD-NATURALISTS’ CLUB, O
, Ottawa, Canada
VOL. LY, No. 8
THE HEIDNATUT
NATURALIST.
published by the ottawa Field-Naturalists’ Club.
CONTEN TS.
: Past
4, Title Page tor Vol 1X, (TAWA NATUR ALIST oc ee aie hes nee 1
3, Patron, Gouneil, Committees and Leaders of The Ottawa Field ‘Naturalists’ Club for 1895-1806, 3
3, Last of Ordinary and Corresponding Members 3! Urn: Pine (nee ts
4, Calourless Biood in Amimals—Prot, Bf, B, Prince, pa, FLS T
& The Rensselaer Grit Plateau RW. Ells, LL.D.) PRS... 7 oe Sale a
8, The Relation of the Atmosphere to Agriculture—F- T. Shutt, MA 5 pa We Beer 12
7. Announcement, Royal Society of Canada ee EE 4
g, Annual Report of Council for 1AGL-1895 1
g, Treasurers Statement -- eae ee aie an ae w
4p, Notes, Review? and ‘Comments, Geategy, Conehalegys Ornitholag > 20
2A
yn. Geological Society of Americt 2-7
AR. Faitorial Note 2 oS
i ere,
OTTAWA, CANADA.
PRINTED BY PAYNTER & ApsoTt,
48 RipEad SrReEr.
Enteral cad the Oltaea Bost Office os secund last matter
1S.
Entered at the ee NOVEMBER 30th, 1941
wa Post Office as cera cl
-class matter
Centennial Year
Volume 93, Number 2
NOVEMBER, 1941
April-J une 1979
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural
heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse
information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving,
maintaining or restoring environments of high quality for living things.
The Members of Council are listed on the inside back cover.
The Canadian Field-Naturalist
The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas
expressed in this journal, however, are private and do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or
any other agency.
Editor: Lorraine C. Smith
Assistant to the Editor: Donald A. Smith Book Review Editor: J. Wilson Eedy
Associate Editors
C3D» Bird A. J. Erskine David P. Scott
E. L. Bousfield Charles Jonkel Stephen M. Smith
Francis R. Cook Charles J. Krebs Robert E. Wrigley
George H. La Roi
Copy Editor: Marilyn D. Dadswell Chairman, Publications Committee: J. K. Strang
Production Manager: Pauline A. Smith Business Manager: W. J. Cody
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The Canadian Field-Naturalist
Volume 93, Number 2. April-June 1979
Our Responsibilities as Field-Naturalists and Biologists
The important message in the guest editorial by Yorke Edwards (Biology — the unknown science?
Canadian Field-Naturalist 93(1): 6-9, 1979) is that the world needs more generalists in science and
that workers in biology must communicate better. My comments here are offered not only as a
reinforcement of his views but also to point out the responsibilities of field-naturalists and biologists
regarding the quality, accuracy, and comprehensiveness of their studies.
Although for years biologists have been isolated from social and political changes, increasing
diaiogue between scientists and the general public and governments 1s taking place. Not all of this is
productive. Nevertheless society is becoming more aware that many of the world’s troubles are
concerned with changes in normal biological processes and ecosystems and that the nature of
environmental problems is complex. In order to formulate policies, decision-makers (normally
situated at the government level) must assess the opinions, often conflicting ones, of scientists. Many
scientists find it difficult to interpret science to the layman. Moreover, with increasing specialization
some scientists can convey information only on limited topics in their own field of interest and are
often unable to comprehend the total picture. Therefore, we need more competent generalists to
integrate and interpret scientific data and to communicate science to society. As Yorke Edwards
wrote, “We need a new morality in the sciences, resulting in major efforts to make science and its
influence on Earth understandable.”
Because the role of scientists in society is becoming more visible, I think it is germane to point out
some of the responsibilities that we should be aware of, but often are not. In particular we should
recognize our limitations, strive for accuracy, and maintain credibility.
It is important, however, not only that we recognize our own scientific limitations but that we do
not exceed them. Unfortunately some biologists extend their studies, interpretations, and
conclusions beyond their level of expertise and competence. Sometimes their conclusions are based
on results obtained using faulty methods and are thus incorrectly attributed to environmental
variation. Sometimes their opinions, whether right or wrong, have been accepted as facts. How does
the decision-maker know whose opinion to respect? This is a major problem which has become even
more complicated by the mushrooming number of biological consulting firms, some excellent but
others abysmally bad. How often has the perhaps glossy, long-winded, eye-appealing final report of
the contractor been valued rather than whether the work on which it is based was scientifically sound?
How often will firms or individuals take on work that is beyond their qualifications? We know that
some environmental consulting firms at least try to make up for their deficiencies in qualified and
competent staff by contacting and “picking the brains” of acknowledged experts (often without any
financial or other compensation or acknowledgment to the experts for their time and knowledge)
whereas others make no attempt to improve the quality of their studies. Some contractors do not
have the ability or inclination to do a proper literature search whereas others may do a search, then
write a research paper under the pretence that their report contains original work.
Biologists or field-naturalists should not be considered specialists outside their own specialty.
Environmental problems in particular are very complex, and what may appear on the surface to be
simple almost always has complicated interrelated ecological aspects. The tendency to look at one
particular aspect of a specific problem without regard for the broader view has sometimes resulted in
113
114 THE CANADIAN FIELD-NATURALIST Vol. 93
disastrous decisions. It is imperative, therefore, that the total involved system be considered if wise
decisions are to be made. But so often this doesn’t happen. Too few are able to see, let alone
comprehend, the total aspect of biological problems and the complicated interrelationships that
exist in nature. Generalists who approach interdisciplinary studies with open eyes and an open mind
are needed to understand and to interpret the basic workings of complex ecosystems, 1.e., to put it all
together. But they too must know their limitations: they cannot be masters of all disciplines;
therefore, they will need to consult specialists on specific aspects of particular problems. Although the
popular interdisciplinary method of teamwork is commendable, often it does not fill the need fora
broad approach. D. B. O. Savile pointed out some years ago (Unity from diversity in biological
research. Transactions of the Royal Society of Canada 4(4): 245-251, 1966) that often each team
member sees only his or her aspect of the problem. He elaborated further as follows: “For the clearest
and fullest picture of any biological phenomenon, it is helpful if the interdisciplinary approach of the
team is complemented by the multidisciplinary approach of the individual.”
We have a responsibility to strive for accuracy. Sometimes incorrect descriptions, identifications,
computations, etc., appear in reports. It is easier to perpetuate an error than to correct one. Studies
should be carried out as thoroughly and competently as possible with careful regard to repeatability,
the basis of the scientific method. Moreover, if statements can be misinterpreted, are ambiguous, or
are otherwise unclear, they should be rewritten.
Often there is considerable pressure to do a study and to prepare the report quickly without
particular regard for accuracy and dependability, and consequently too much may be read from
limited data. Researchers should be obliged to state the limitations of their methods, data, and
conclusions so that there will be less distrust and fewer set-backs but rather an advancement of
knowledge.
We must endeavor to maintain our credibility and should encourage other biologists and field-
naturalists to do so too. This is particularly important when the problems and issues that arise are
emotional ones. Interpretations, recommendations, and conclusions should, as far as possible, be
based on good sound scientific data. Speculation should be limited to what can be reasonably well
supported. We mustn’t compare apples and oranges. Thus the results obtained on one species cannot
be directly extrapolated to another species.
Our credibility is dependent on the accuracy of our statements, on recognizing the limitations of
the data, and on the limitations of our expertise. Obviously if errors are discovered in one part of our
work, the results from an entire study will be suspect. For example at a recent naturalists’ meeting, a
petition for signatures was presented. Although this petition had an admirable conservation measure
as an objective (it was against the legalization of the taking of certain birds of prey), alas the reasoning
accompanying it was false. It was alleged that these particular raptors should be saved because as
rodent-eaters they are beneficial to farmers. When it was pointed out that these particular birds prey
mainly on birds, not rodents, the petition was not withdrawn for this error to be corrected. In such
cases, where we make false statements, opponents will dwell on them, and a good cause may be lost.
A rather copious volume of “gray” scientific “literature” or pseudoliterature has appeared in
recent years. Some of this has resulted from the many contracts let for baseline surveys and
preparation of environmental impact statements. The greatest shortcoming of the duly written final
reports is that they are not generally subjected to peer review as are research papers published in the
primary literature. Frequently errors of commission and omission are put into such records and are
perpetuated. Considerable time and effort are required to correct these. Indeed some are never
corrected. The solution to this problem is not easy. Because they are not subjected to rigorous
(or any in some cases) peer review, there is a tremendous variability in the reports in the gray liter-
ature: some reports are very superficial but others are excellent in-depth studies. Better evaluation
of these reports is needed.
Floral and faunal surveys of many areas, especially national and provincial parks and areas
designated by the International Biological Programme, are indeed valuable. But how can a
judgment be made on whether the contractors are really trained and reliable? One partial answer is
1979 SMITH: EDITORIAL WS)
to be sure voucher specimens are kept, because these at least can be checked for correct
identification. We know of at least one government contract where the employees of a consulting
firm had misidentified and reported incorrectly ona large proportion of the specimens. It is cases like
this that when discovered cause credibility to be lost. But how many go undetected? Many biologists
and field-naturalists are concerned about these quick and superficial surveys, with the
overconfidence of some investigators, and in the final analysis with credibility.
Other scientists are also worried about these problems. In his editorial in the first issue of a new
journal Marine Environmental Research (July 1978) Eric Cowell wrote: “Nowhere has departure
from scientific integrity been more evident than in the field of environmental research. .. . There is
much to learn about the function of our ecosystem and man’s role in the management or mismanage-
ment of it. We need clearly defined objectives, impeccable data and analyses and interpretation
conducted with integrity.”
My plea to field-naturalists and biologists is to follow the old adage “if it’s worth doing, it is worth
doing well.” Of course it takes time and effort to do good research, whether it is in the field, in the
laboratory or museum, or in the library. Check for accuracy, maintain scientific integrity, exercise
care in the interpretation of data, and recognize the limitations. Seek the comments of specialist
colleagues but look to generalists to give the total picture. Only if we recognize and carry out these
responsibilities can we hope to maintain quality and standards and to contribute positively to society
as responsible citizens.
LORRAINE C. SMITH
Editor
Natural Fires as an Index of Paleoclimate
J. TERASMAE! and N. C. WEEKS?
'Department of Geological Sciences
2Department of Biological Sciences, Brock University, St. Catharines, Ontario L2S 3A]
Terasmae J. and N. C. Weeks. 1979. Natural fires as an index of paleoclimate. Canadian Field-Naturalist 93(2): 116-125.
The charcoal abundance and frequency of occurrence varies stratigraphically in postglacial lake sediments, and this
information can be used together with palynological and sedimentological data for a reconstruction of paleoclimatic
conditions. The gelatin-coated slides method can be used for continuous sampling of lake sediment cores to determine the
presence of charcoal particles. Changes in the charcoal profile have been related to changes in the sequence of fossil pollen,
specifically that of pine pollen, and from the observed relationship a climatic control of forest fires is inferred. Forest fires
have occurred naturally in the Great Lakes — St. Lawrence and Boreal forest regions during the past 9000 yr, and the fire
frequency almost doubled during the Pine Pollen Zone about 7000 to 4000 yr ago in the southern Boreal forest. The mean fire
frequency in the Lac Louis area was one fire every 95 to 100 yr, but during the period characterized by a pine pollen maximum,
7280 + 240 yr BP (GSC-1481) to 4260 + 240 yr BP (GSC-1491), the fire frequency increased to one fire every 48 to 56 yr.
Key Words: forest fires, palynology, paleolimnology, paleoclimate.
Natural fires, as distinct from man-induced
ones, have been characteristic of terrestrial
ecosystems probably for many millions of years.
The prerequisites are (1) a readily combustible
‘fuel’ source (vegetation) and (2) an igniting
mechanism. The first of these is_ satisfied
whenever and wherever a supply of dry vege-
tation is available, and both lightning and
volcanism have been present as igniting mecha-
nisms throughout geological time; however, the
frequency of coincident fuel, ignition, and fire
has varied through time.
The fact that numerous and very extensive
forest fires, as well as grassland and also tundra
fires, coincide with general or seasonal dry
weather episodes (drought) appears to be com-
monly accepted (Rowe et al. 1974, 1975).
Meteorological conditions leading to such epi-
sodes of drought are caused by lack of precipi-
tation, relatively high summer temperature,
extended periods of sunny weather, and medium-
to high-velocity surface wind. A combination of
these meteorological variables will intensify the
development of drought conditions and, hence,
increase the susceptibility of vegetation to fire.
The linkage between climatic conditions and the
frequency and areal extent of natural fires in
vegetation seems to be well established. There-
fore we assume that if the frequency of natural
fires can be determined by studies of sediment
records preserved in lakes and peat bogs, it will
116
be possible to interpret this information in terms
of past climatic conditions and to use the time
frequency of natural fires as an indicator or
index of paleoclimate.
There has been also an increasing interest in.
the place of natural fire in the environment over
the past few years, paralleled by a growing
perception of fire as a natural and necessary
environmental factor rather than an unnatural
“disturbance” that must be controlled and
eliminated whenever and wherever possible.
The presence of charcoal fragments is the
primary evidence that natural fires leave in the
sedimentary record, as indicated already by
other studies (e.g., Heinselman 1971, 1973;
Swain 1973: Rowe and Scotter 1973).
The purpose of this study is to investigate the
methods used for extracting fire frequency
records from lake sediments, and the criteria
for selecting appropriate sites for this kind of
study.
Lake sediments constitute a natural deposi-
tory of environmental information that accumu-
lates year by year throughout the life-span of a
lake. These sediments contain inorganic (sand,
silt, clay) particles and organic detritus (pollen,
spores, diatoms, cladocera, various other algae
and fungi, identifiable plant tissue fragments,
and charcoal), as well as chemical residues,
precipitates, break-down products, and stable
compounds. All these components collectively
STS
reflect the environmental conditions within the
lake as well as in the surrounding area. A
stratigraphic study of these sediment com-
ponents can establish a time series of ecological
events from which paleoecological and environ-
mental changes can be inferred on the basis of
modern analogues (studies of recent limno-
logical-environmental relationships).
The study was designed to (1) explore the
stratigraphic occurrence of charcoal in sedi-
ments of selected lakes for the purpose of
establishing a charcoal frequency record for
postglacial (Holocene) time, (2) infer from this
record the frequency of natural fires (forest
fires), and (3) relate this information to the
paleoenvironmental changes (including climate
and vegetation) that have been established for
the same time interval on the basis of palyno-
logical studies of the same sediment cores.
It was assumed that charcoal particles, origin-
ating from forest fires, are transported to the
lake by atmospheric turbulence (wind), possibly
by surface water runoff, and direct falling of
charred plant matter into the lake. Charcoal
influx, contributed by all means of transport,
would be further dispersed in the lake by surface
currents and eventually settles to the bottom
where it is incorporated in the accumulating
sediment as an identifiable record of the fire.
Location and Description of the Study Sites
The study sites were four lakes (Figure 1)
selected because of availability of background,
palynological studies, radiocarbon dating, map-
ping of surficial deposits, and the knowledge of
postglacial history of the region. Furthermore,
palynological studies of lake sediments and peat
deposits made at numerous sites throughout
Ontario by the senior author over a period of
some 20 years (Terasmae 1973) had demon-
strated common occurrence of charcoal particles
in many palynological preparations. There
seemed to be an apparent lack of published
information from southern Ontario on charcoal
occurrence in lake sediments and its relation-
ship to the postglacial history of natural fires.
Three lakes are in the Great Lakes — St.
Lawrence Forest region (Found Lake,
45°30'65’N, 78°30'80”W; Perch Lake, 46°02’N,
77°21’W; and Boulter Lake, 46°09’15’N,
79°02’W) whereas Lac Louis (47°15’15’N,
TERASMAE AND WEEKS: FIRES AS AN INDEX OF PALEOCLIMATE
Ly)
79°07’W) is in the southern part of the Boreal
Forest region. The Great Lakes — St. Lawrence
Forest is characterized by a mixture of
deciduous hardwood tree species and some
coniferous tree species, but coniferous tree
species attain dominance in the Boreal Forest
(Rowe 1959). Some descriptive data on the four
lakes and their sediments are summarized in
Table 1.
All four lakes are different in terms of their
geological characteristics. Found Lake is a
bedrock basin, surrounded by steep rock slopes,
and characterized by only minor inflow of
surface and ground water. It is oligotrophic and
has a low sedimentation rate (0.41 mm per year).
Perch Lake is a depression in surficial deposits
(mostly sand) and only a minor part of the shore
is bordered by bedrock. It receives some ground-
water and surface-water inflow through sur-
rounding swamp and bog vegetation that filters
out particulate matter, and a small creek flows
out of Perch Lake to Ottawa River. It is about
8-10 km from the large sand dune area of the
Petawawa delta. The lake is eutrophic and has
an average sedimentation rate of 0.61 mm per
year.
Boulter Lake occupies an ice block depression
(a “kettle hole”) in glacial outwash and esker
deposits. It has no bedrock shoreline and
probably receives some inflow of groundwater.
There are no streams flowing into or out of the
lake, and it is eutrophic; average sedimentation
rate is 0.66 mm per year.
Lac Louis (elevation 300 m) is located just
below the highest shoreline (305 m) of glacial
Lake Barlow-Ojibway in a sand and gravel
deposit of an esker complex (Vincent 1973), and
it is probably a “kettle” lake. It receives some
inflow of groundwater and a small stream flows
out of it. It is surrounded by Black Spruce ( Picea
mariana) forest and muskeg, with some Trem-
bling Aspen (Populus tremuloides) and White
Birch (Betula papyrifera). The lake is eutrophic;
average sedimentation rate 1s 0.59 mm per year.
The lakes share a common characteristic,
being essentially closed basins, without any
major inflowing streams and with only minor
outflow. Furthermore, all lakes are surrounded
(except where bedrock forms the shore) by
wetland vegetation (bog and swamp) that effec-
tively prevents input of particulate matter by
118 THE CANADIAN FIELD-NATURALIST Vol. 93
1 Found Lake
2 Lac Louis
3 Perch Lake
4 Boulter Lake
I
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ONTARIO QiViEBEG
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Fo&smill \
o ALGONQUIN \ 3X
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FIGURE |. Index map, showing the location of study sites. Found Lake (site 1), Perch Lake (site 3), and Boulter Lake
(site 4) are in the Great Lakes — St. Lawrence forest region, and Lac Louis (site 2) is in the southern part of the
Boreal forest region.
TABLE |—Some measurements and comparative data on the four lakes studies
Found Lake Perch Lake Boulter Lake Lac Louis
Elevation (m) 460 156 389 300
Maximum depth
of water (m) 34 355) 8.5 7.6
Thickness of organic
sediment cored (cm) 430 600 780 540
Surface area (km2) 0.32 0.45 0.25 0.10
Age of oldest organic 10400 + 300 9830 + 250 11800 = 400 9090 + 240
sediment (years BP) (I-7782) (GSC-1516) (GSC-1363) (GSC-1432)
Average sedimentation
rate (mm per year) 0.41 0.61 0.66 0.59
1979
surface water runoff. It is assumed, therefore,
that charcoal particles found in the lake sedi-
ment have been transported to the lakes pri-
marily by atmospheric dispersal. The only
exception would be charcoal input by water
runoff from the small area of bedrock slopes
adjacent to the lake. This assumption is sup-
ported by the observation that many charcoal
particles in the lake sediment are in the same size
range as pollen (about 20 to 150 um) that is
transported to the lakes by atmospheric disper-
sal. All four lakes are also characterized by very
small watersheds, effectively restricted to the
surrounding slopes extending only 100-200 m
away from the lake. Only Perch Lake has a
larger catchment area that is occupied by
wetland vegetation.
Methods
The lake sediment cores were taken with
piston corers; the Brown sampler was used for
very soft sediment at and near the water-
sediment interface, and the Livingstone sampler
for the remainder of the organic and inorganic
sediments cored (Mott 1966), after a recon-
naissance of lake basin morphometry (depth
sounding) and the surrounding landscape had
been made. Palynological subsampling of sedi-
ment cores, laboratory preparations, and micro-
scopic examination of slides followed well
established procedures (Faegri and Iversen 1975;
Kummel and Raup 1965).
The study of charcoal presence and abun-
dance was carried out in two different ways. In
one case the palynological slides were used for
counting of charcoal particles. The principal
problem in this case is that samples for a
palynological study (0.5 to | cm3) are normally
taken at 10-cm (or sometimes 5-cm) intervals
from the sediment core, and each sample covers
sediment deposition during 10-20 yr, depending
on sedimentation rate. This method of sampling
obviously does not provide any information for
the time intervals (about 100-200 yr) repre-
sented by sediment deposition between the
sampled intervals; therefore, it is possible that a
significant part of the record of forest fires may
be missed.
To overcome this problem in the charcoal
study, Weeks developed another preparation
method, based on a technique used for sedi-
TERASMAE AND WEEKS: FIRES AS AN INDEX OF PALEOCLIMATE 119
mentary grain-size analysis by Perrie and Peach
(1973) and Peach and Perrie (1975). The method
involves two principal steps: preparation of
gelatin-coated slides, and transfer of sediment
onto the slide.
The slides were prepared by first dissolving 1.5
teaspoons “Knox Gelatin” in 300 mL distilled
water (the water-gelatin mixture was warmed in
a water bath). Clean glass slides were then
dipped in the gelatin coating solution (except the
frosted ends, used for labelling). The excess
gelatin was allowed to run off the slides, which
were then placed in a dust-free cabinet to dry for
a maximum of 6 h before storage in a clean slide
box.
The second step began with preparation of the
sediment core for sampling. The sediment
surface was carefully scraped clean of possible
contamination with a spatula, and this surface
was made smooth so that the full gelatin-coated
area of the sampling slide came into contact with
the sediment when the slide was touched or
pressed against the core.
It should be noted that the sediment cores,
after they had been extruded from the coring
tubes, were wrapped in plastic and aluminum
foil and stored ina temperature-controlled room
at 3-5°C to retain the natural moisture content,
a normal practice for palynological studies.
The sampling was carried out by pressing the
gelatin-coated slide lightly to the sediment core:
a thin layer of moist sediment sticks to the dry
gelatin surface and a “fingerprint” of particles is.
lifted off. The amount of pressure on the slide,
required for producing a good “fingerprint,”
depends on sediment characteristics and mois- °
ture content and this is determined experi-
mentally by examining the slides under a micro-
scope to make sure that the optimum amount of
sediment is picked up by the slide. Too much
sediment on the slide makes counting of char-
coal particles difficult; too little sediment re-
quires the counting of these particles on several
slides, which is rather time-consuming. It is
expected that further experimentation will lead
to a more specific description of this aspect of the
method. This sampling method covers the total
length of the core without any gaps between
samples.
Cover slips were placed on the slides and a
small amount of xylene was added to the edge of
120
the cover slip. The xylene dispersed throughout
the preparation and rendered most of the
organic detritus matrix translucent, whereas
charcoal particles, pollen grains, and diatoms
remained clearly visible. The ‘clearing effect’ of
xylene made the identification and counting of
charcoal particles much easier (Figure 2).
Charcoal particles were identified with the aid
of samples collected from sites of recent forest
fires and prepared according to methods used in
this study. Normally these particles are readily
identifiable because some larger (commonly
wood) fragments are only partially charred and
others still retain characteristics of recognizable
plant tissue structure. Some smaller charcoal
particles are opaque (black) and differ from
mineral grains with respect to fracture, refractive
index, and surface characteristics. The common
mineral grains are, furthermore, mostly trans-
parent and lack cellular structure. The problem
of small pyrite grains that occur in some lake
sediments (even inside of pollen grains), how-
ever, requires further investigation.
The charcoal particles were counted in three
different ways. In the first case, charcoal was
counted together with pollen on slides prepared
for palynological study. This technique provides
a direct comparison of charcoal abundance with
palynological data in the same preparation.
In the second case the charcoal particles were
counted on the gelatin-coated slides. The pur-
pose of this technique was to relate charcoal
abundance to sediment stratigraphy because
each slide covers about 5cm of sediment
deposition.
In the third case, an attempt was made to use
the Quantimet 720 (a TV image-analyzing
computer) for counting of charcoal particles on
the gelatin-coated slides. The advantage of this
technique is that the instrument can be set to
count particles in predetermined size classes,
as described by Peach and Perrie (1975) in their
study of varved glacial lake sediments. There
was some difficulty, however, in having the
Quantimet distinguish charcoal particles from
other similar particles, and further experiments
will have to be made to resolve this problem.
For sediment cores from Lac Louis, Found
Lake, and Boulter Lake the charcoal occurrence
and relative abundance were tabulated on the
basis of data obtained from continuous
THE CANADIAN FIELD-NATURALIST
Vol. 93
sampling by the gelatin-coated slides method.
For Perch Lake the charcoal data were obtained
by examination of the palynological slides and
palynological data were adopted from available
pollen diagrams (Boyko-Diakonow and
Terasmae 1975). The charcoal data for the three
other lakes (Lac Louis, Boulter Lake, and
Found Lake) were obtained from the same
sediment cores that had been used for palyno-
logical study (Vincent 1973; R. J. Mott, personal
communication; M. Boyko-Diakonow,
personal communication), and a direct com-
parison of both kinds of data was again possible.
In Figure 3 the abundance of charcoal was
calculated in the same manner as the relative
pollen percentages of the different tree species;
1.e., the total number of pine pollen grains, for
example, is expressed in the pollen diagram as a
percentage of the total tree pollen count in a
particular preparation (sample).
The determination of sedimentation rate was
considered an important aspect of the study of
stratigraphic dispersal of charcoal in lacustrine
deposits because the estimated (or assumed)
sedimentation rate provides the time scale for
chronostratigraphy of charcoal occurrence and
abundance, unless some other means of time
calibration is available (for example, when the
sediment is annually laminated, or varved).
Radiocarbon dates were available for all sedi-
ment cores studied and these dates were used to
calculate the approximate sedimentation rates
that are shown in Table | for each of the lakes
investigated.
All tabulated data used as the basis of this
paper are included in an unpublished report by
Weeks (B.Sc. thesis, available from Department
of Biological Sciences, Brock University).
Results and Discussion
Our observations indicate that charcoal does
not occur in distinct layers with well defined
boundaries in the lake sediments studied. Al-
though the abundance of charcoal particles
changes stratigraphically, the boundaries of
each occurrence of these particles are diffuse,
implying that at least some mixing of sediment
has taken place during deposition and that
charcoal dispersal (in decreasing amounts) may
have occurred during several years after the fires
that are recorded in the lake sediment.
1979 TERASMAE AND WEEKS: FIRES AS AN INDEX OF PALEOCLIMATE 12]
FIGURE 2. Photomicrographs of gelatin-coated slides from Boulter Lake sediment core: (A) untreated, (B) the same slide
after clearing with xylene; and Found Lake: (C) untreated, (D) cleared with xylene. Arrows — charcoal particles;
black triangle — pine pollen grain.
122 THE CANADIAN FIELD-NATURALIST Vol. 93
CUPRESSINE AE
DEPTH (cm)
aa a a Sa a I pr arene fener rel emcee fee pon fa
IDO OR ZORS SONG 10 20 30 40 50 60 70 80 0 lOR ZO SOLOREOIO Om co
98307250 (GSC !516) POLLEN PERCENTAGES W@ CHARCOAL
POLLEN
a
a
>)
Ee
WwW
{ea)
DEPTH (em)
GSC 1481
7280+250
POLLEN PERCENTAGES
FIGURE 3. Abbreviated pollen diagrams from Perch Lake (A) and Lac Louis (B). The black areas indicate occurrence
and abundance of charcoal particles, based on study of palynological slides (Perch Lake) and gelatin-coated slides
(continuous sampling) from the Lac Louis sediment core.
1979
The diffuse nature of the charcoal layers in lake
sediments has not been explained adequately by
this study and the subject of modern charcoal
dispersal and deposition in lakes requires more
investigation.
In the Perch Lake study all palynological
slides contained some charcoal, but in view of
the large sampling interval (about 10 cm) as
compared with the annual sedimentation rate
(0.61 mm) it is not possible to be certain of the
actual stratigraphic frequency of charcoal oc-
currences. Quite certainly many _ charcoal
occurrences are not recorded by this type of
study and the question of how much information
was missed in the Perch Lake study will be
investigated further.
The Found Lake sediment core contained
relatively little charcoal, both in terms of
stratigraphic occurrence and abundance. On the
other hand, the part of Boulter Lake sediment
core (640-750 cm) that was examined in this
study (because palynological data were available
only for the basal 2 m of this core) contained
abundant charcoal at frequent intervals.
The Lac Louis sediment core showed a rather
high frequency of charcoal occurrence with a
non-uniform stratigraphic distribution and
varying abundance. A pollen diagram for Lac
Louis has been published by Vincent (1973).
The sparseness of charcoal in the Found Lake
sediment core can be attributed to at least two
possible causes. One is the site susceptibility to
fire, because the forest of the Found Lake area is
dominated by hardwoods, mainly maple (Acer),
beech (Fagus), and birch ( Betula). The second is
related to the seasonal frequency distribution of
thunderstorm activity. According to Chapman
(1952) some thunderstorms in this area coincide
with the period of fire susceptibility in the
hardwood forest, but the greatest thunderstorm
activity occurs in June, July, and August when
the trees are in full leaf. Autumn fire hazard is
reduced by the increased precipitation during
that season, and Chapman refers to this north-
ern hardwood forest as being the most nearly
immune to fire of all forest types.
These two causes may explain the paucity of
charcoal in the Found Lake sediments during
the last 7000 to 8000 yr but this reasoning
becomes much weaker for the earlier postglacial
time, when, on the basis of palynological
TERASMAE AND WEEKS: FIRES AS AN INDEX OF PALEOCLIMATE
8
evidence, the forest in this area contained more
pine and fewer hardwood species some 8000 to
10 000 yr ago, and the forest was dominated by
spruce prior to 10000 yr ago. Therefore, it is
unlikely that vegetation alone was the factor in
fire resistance in early postglacial time, although
it may be now. Probably other factors, including
topography, microrelief, and soil moisture are
also important, and Found Lake may not be a
good site for carrying out forest fire frequency
studies.
Charcoal particles were present throughout
the Lac Louis sediment core, and the charcoal
profile as shown in Figure 3 was based on the
continuous sampling procedure, using gelatin-
coated slides. Unfortunately the top and bottom
4 m of sediment core was not available for this
study, and hence no charcoal profile is shown for
these parts of the core in Figure 3.
Interpolation of the radiocarbon dates for Lac
Louis (reported by Vincent 1973) gives an age of
about 8100 yr BP at the 480-cm depth of
sediment. Within the 480 cm, there are 84
charcoal occurrences. If we assume that each is
related to a separate fire event, there is one fire
every 95 to 100 yr. This is probably an inaccurate
estimate, however, because the distribution of
charcoal is stratigraphically non-uniform, and a
simple expression of mean frequency fails to
express the fluctuations of fire frequency
throughout the sediment core. For example, fire
frequency in the pine zone (Pollen Zones 3 and 4)
of the Lac Louis sediment core (Figure 3) is one
fire every 48 to 56 yr — at least double the mean
fire frequency for the whole core.
Although only a part of the Boulter Lake ©
sediment core was examined for charcoal
(640-750 cm), this part of the core contains the
pine pollen zone (R. J. Mott, personal com-
munication) and both a high frequency and
abundance of charcoal. It seems that in Lac
Louis, and probably in Boulter Lake, the high
frequency and abundance of charcoal coincides
with high percentages of pine pollen (the Pine
Pollen Zone).
For Perch Lake, the fire frequency, based on
examination of pollen slides, is calculated to be
one fire every 140 to 150 yr. This isa much lower
fire frequency than one would expect, given the
long history of fire which is evident from
palynological data and an environment char-
124
acterized by sand dunes of the Petawawa delta
and pine-dominated vegetation. This value also
conflicts with the findings of Cwynar (1975) ata
similar site, about 50 km to the west where the
mean fire frequency in Greenleaf Lake area over
the past 1200 yr is one fire every 83 yr. It seems
reasonable to conclude that a number of
charcoal occurrences have been missed owing to
the distance between !0-cm sampling intervals.
A study of Lake of the Clouds, Minnesota,
relates forest vegetation succession over the past
10 000 yr to climatic change, and charcoal data
to the changes in forest vegetation (Swain 1973).
Swain reports an increase in fire frequency after
9500 yr BP, which coincides with a pine pollen
peak and a climatic warming trend (the hypsi-
thermal) which reached an optimum about
7000-6000 yr BP. Consistent high charcoal
abundance occurred from 9000 to 6000 yr BP.
Then a period of climatic cooling began, spruce
and alder increased, White Pine ( Pinus strobus)
decreased, and charcoal abundance became
irregular from 6000 to 3000 yr BP. From 3000 to
1200 yr BP, a decrease in charcoal occurrence is
associated with a similar decline in White Pine.
The similarity between the pattern of climate,
vegetation, and fire occurrence described by
Swain at Lake of the Clouds and that reported
by Vincent (1973) and this study at Lac Louis is
rather striking. The same stratigraphic pattern
of charcoal distribution seems to occur at both
sites, but the Lac Louis dates are somewhat
younger, possibly because of the difference in the
time of deglaciation. The Lake of the Clouds
area was deglaciated about 14 500 yr BP, but Lac
Louis did not become ice free until about 10 500
yr ago. This implies that vegetation migrated
northward with the receding ice and, therefore,
radiocarbon dates from Lac Louis are about
7280 + 240 yr BP (GSC-1481) for the beginning
of Pollen Zone 3 and 4260+240 yr BP
(GSC-1491) for the end of this zone (the Pine
Pollen Zone), compared to 9000 and 6000 yr BP
for the Minnesota site.
Both Lac Louis and Lake of the Clouds lie
within the Great Lakes -St. Lawrence and
Boreal Forest transition zone. There seems to be
good potential for paleoclimatic reconstruction
on the basis of charcoal occurrence in, and pollen
analysis of lake sediment cores within, this zone.
THE CANADIAN FIELD-NATURALIST
Vol. 93
The gelatin-coated slide technique for charcoal
analysis could be useful in such regional paleo-
climatic studies.
The results of the present study suggest that it
will be possible to work out a postglacial forest
fire frequency sequence for at least some
regions, such as the deciduous forest and the
boreal forest when study sites are selected with
appropriate care. Geological site characteristics
(topography, types of rocks and surficial de-
posits, surface and ground water regime, glacial
history and landscape features such as deltas,
outwash plains, eskers, etc.) should be con-
sidered with respect to fire susceptibility of the
vegetation that grows on these different sites.
For example, the Found Lake area seems to be
rather resistant to forest fires: the Perch Lake
area, in contrast, appears to have been especially
susceptible to forest fires because of geological
characteristics of the landscape that at least
locally control the composition of forest. There-
fore, these areas are less suitable for forest fire
frequency studies than the Lac Louis area, or
the Lake of the Clouds area in Minnesota.
It appears possible that a fire frequency index
can be worked out for Holocene lake sediment
sequences and related to paleoclimatic condi-
tions that have been inferred, in part, from other
kinds of evidence such as palynology and the
physical and chemical sediment characteristics.
Acknowledgments
We gratefully acknowledge the assistance of
R. J. Mott of the Geological Survey of Canada
who generously provided some sediment core
material and some palynological data. We thank
J. Simpson (superintendant), D. Brunton, and
D. Strickland of the Algonquin Provincial Park
for their help with the Found Lake study and for
permission to use data from their files, including
a pollen diagram compiled for Found Lake by
J.H. McAndrews of the Royal Ontario
Museum. Special thanks are extended to George
H. La Roi for helpful criticism and many
suggestions that were invaluable for the
improvement of this paper. Partial financial
assistance for this study was provided by the
National Research Council of Canada
(operating grant A5545 to J. Terasmae).
1979
Literature Cited
Boyko-Diakonow, M. and J. Terasmae. 1975. Palynology
of Holocene sediments in Perch Lake, Chalk River,
Ontario. /n Hydrological studies on a small basin on the
Canadian Shield. Edited by P. J. Barry. Atomic Energy
of Canada, Ltd., Report AECL-5041-I, Chalk River,
Ontario. pp. 189-220.
Chapman, H.H. 1952. The place of fire in the ecology of
pines. Bartonia 26: 39-44.
Cwynar, L. 1975. The fire history of Barron Township,
Algonquin Park. M.Sc. thesis, Department of Botany,
University of Toronto, Toronto, Ontario.
Faegri, K. and J. Iversen. 1975. Textbook of pollen
analysis. 3rd edition. Blackwell Scientific Publications,
Oxford. 295 pp.
Heinselman, M. L. 1971. The natural role of fire in North-
ern Conifer Forests. /n Proceedings Fire in the northern
environment —a symposium. Edited by C. W. Slaughter,
R. J. Barney, and G.M. Hansen. Pacific Northwest
Forest and Range Experiment Station, Portland, Oregon.
pp. 61-72.
Heinselman, M. L. 1973. Fire in the virgin forests of the
boundary waters Canoe Area, Minnesota. Quaternary
Research 3: 329-382.
Kummel, B. and D. M. Raup. (Editors). 1965. Handbook
of paleontological techniques. W. H. Freeman and Com-
pany, San Francisco. 852 pp.
Mott, R. J. 1966. Quaternary palynological sampling tech-
niques of the Geological Survey of Canada. Geological
Survey of Canada, Paper 66-41, Ottawa. 24 pp.
Peach, P. A.and L. A. Perrie. 1975. Grain-size distribution
within glacial varves. Geology 3(1): 43-46.
TERASMAE AND WEEKS: FIRES AS AN INDEX OF PALEOCLIMATE
WZ)
Perrie, L. A.and P. A. Peach. 1973. Gelatin-coated micro-
scope slides in sedimentary size analysis. Journal of
Sedimentary Petrology 43(4): 1174-1175.
Rowe, J.S. 1959. Forest regions of Canada. Department
of Northern Affairs and National Resources,. Forestry
Branch, Bulletin 123. 71 pp.
Rowe, J.S. and G. W. Scotter. 1973. Fire in the boreal
forest. Quaternary Research 3: 444—464.
Rowe, J.S., J. L. Bergsteinsson, G. A. Padbury, and
R. Hermesh. 1974. Fire studies inthe Mackenzie Valley.
Department of Indian Affairs and Northern Develop-
ment, Northern Economic Development Branch, ALUR
Program, Report ALUR 73-74-61. 123 pp.
Rowe, J. S., D. Spittlehouse, E. Johnson, and M. Jasieniuk.
1975. Fire studies in the Upper Mackenzie Valley and
adjacent Precambrian Uplands. Department of Indian
Affairs and Northern Development, Northern Economic
Development Branch, ALUR Program, Report ALUR
74-75-61. 128 pp.
Swain, A. M. 1973. A history of fire and vegetation in
N.E. Minnesota as recorded in lake sediments. Quaternary
Research 3: 383-396.
Terasmae, J. 1973. Notes on late Wisconsin and early
Holocene history of vegetation in Canada. Arctic and
Alpine Research 5(3) Part 1: 201-222.
Vincent, J. S. 1973. A palynological study for the Little
Clay Belt, northwestern Quebec. Naturaliste Canadien
100: 59-70.
Received 10 November 1976
Revised and resubmitted 27 June 1978
Accepted 31 October 1978
Virgin Douglas Fir Forest on
Saturna Island, British Columbia
THOMAS P. SULLIVAN!
Department of Botany, University of British Columbia, Vancouver, British Columbia V6T 1W5
'Present Address: Institute of Animal Resource Ecology, University of British Columbia, Vancouver, British Columbia
V6T 1W5
Sullivan, T. P. 1979. Virgin Douglas Fir forest on Saturna Island, British Columbia. Canadian Field-Naturalist 93(2):
126-131.
A survey of the Saturna Island Ecological Reserve provided a description of its virgin Douglas Fir ( Psewdotsuga menziesii)
forest. The vegetation was quantitatively analyzed with respect to structure and composition. The tree stratum dominated the
structure of this reserve, followed secondarily by the low-shrub and bryophyte strata. Psewdotsuga menziesii was the most
important tree species with Tsuga heterophylla (Western Hemlock) and Thuja plicata (Western Red Cedar) probablylimited
by low precipitation and soil texture. Gaultheria shallon (Salal) was the dominant understory species with Eurhynchium
oreganum and Hylocomium splendens dominating the bryophyte stratum.
Key Words: British Columbia, coniferous forests, ecological reserve, forest structure, old-growth forests, Pseudotsuga
menziesii.
Ecological reserves have been established in
British Columbia primarily for scientific
research and related educational purposes per-
taining to our natural environment (Krajina
1973). Included within this proclamation (Eco-
logical Reserves Act 1971) is the conservation of
natural communities over a long period of time
to prevent their alteration by man. The pre-
servation of rare or endangered native plants
and animals in their natural habitat is of
paramount importance in this concept of con-
servation.
The selection of reserves has been based
largely on the previous ecological classification
of the province into biogeoclimatic zones.
Saturna Island and its ecological reserve,
composed of virgin coastal Douglas Fir forest,
are located in the dry subzone of the Coastal
Douglas Fir (CDF) zone (Krayjina 1965, 1969).
An IBP (International Biological Programme)
survey of the area was completed in 1968 by
T.M.C. Taylor and T.C. Brayshaw (summary
sheet of Region |, Area 4 in La Roiet al. 1976).
The objective of this study was to provide a
detailed ecological description of the virgin
Douglas Fir forest on this reserve. This type of
description is essential to the understanding and
management of ecological reserves.
Study Area
Saturna Island, comprising approximately
28 km2, is the most southern Canadian island of
126
the Gulf Islands in the Strait of Georgia. It hes
between 48°46’-48°49’N and 123°2’-123° 13’W.
The Saturna Island Ecological Reserve is
composed of two quartersections and encom-
passes 1.3 km2. It has a range of elevation of
from 150 to 320 m. Figure | shows the interior of
this Douglas Fir forest.
Weather data for this region are lacking. The
study area is characterized by mild temperatures
with prolonged cloudy periods, especially in
winter, and a small range of temperature
seasonally. Fairly wet but mild winters and
warm dry summers are prevalent with a long
frost-free season. Annual total precipitation is
less than 90 cm because of the rain shadow effect
of the Olympic Mountains and higher elevations
on Vancouver Island.
Methods
Vegetation sampling was done during July
and August 1972. The vegetation was analyzed
by strata. The tree, shrub, and high-herb strata
were sampled (restricted random sampling
scheme) by 21 plots, 10 X 20 m each. The low-
herb-dwarf-shrub stratum and bryophyte-lichen
stratum were sampled by 210 subplots, each
1X 1m; 10 were placed systematically within
each large plot.
Data were obtained for diameters of all trees
over 10 cm dbh (diameter at breast height) in the
plots by species; heights of representative trees in
plots using a Blume-Leiss altimeter; density of
SULLIVAN: DOUGLAS FIR FOREST, B.C.
27)
FiGuRE |. Interior of stand of trees representative of the virgin Douglas Fir forest on the Saturna Island Ecological Reserve.
tree saplings (individuals <10cm dbh) by
species; density of tree seedlings in subplots by
species; percent cover estimates for all plant
species in their respective plots and subplots
using the coverage scale of Daubenmire (1959,
1968). A complete plant species list for the
Saturna Island Ecological Reserve is available
from Depository of Unpublished Data, CISTI,
National Research Council of Canada, Ottawa,
Canada KIA O0S2. Vascular plant nomenclature
follows Hitchcock and Cronquist (1973), that of
bryophytes follows Lawton (1971) and Stotler
and Crandall-Stotler (1977), and that of lichens
follows Hale and Culberson (1970).
Results
Importance values, absolute measurements,
and diameter size classes for the tree stratum are
presented in Table 1. The importance value is a
summation of percentages of relative density
+ relative basal area + relative frequency (after
Curtis and McIntosh 1951; Bray and Curtis
1957; Curtis 1959). Pseudotsuga menziesii has
the highest importance value at 190 and the most
consistent distribution of size classes throughout
the stand. The average percent cover values for
trees and saplings are presented in Figure 2.
Douglas Fir trees are located on every plot and
exhibit consistently high cover values through-
out the reserve. This dominant species has a
mean cover value of 52%. Unfortunately, it was
not possible to obtain core samples from the
protected trees on this reserve (regulations
prohibit any disturbance of vegetation on
ecological reserves), and hence there are no
estimates of stand age. Average percent cover
values for the three species in the low-shrub
stratum and the two fern species in the high-herb
stratum are in Figure 2. Gaultheria shallon
(Salal) is distributed throughout most of the
reserve and completely dominates the low-shrub
layer. The average percent cover values for the
most dominant species in the low-herb — dwarf-
shrub and bryophyte strata are presented in
128
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE I—Importance values, absolute measurements, and diameter size classes.for tree species in the Saturna Island Ecological Reserve, based on measurements
in 21 plots, 10 X 20 m each
> 10cm dbh — size class distribution (cm)
density-ha !
Importance Absolute Absolute Absolute Max
value* density basal area frequency <10 cm ht.
Species (0-300) (ha7!) (m2?.ha"!) (%) Seedlings dbh 10- 15- 20- 30- 41- l= 61- (m)
Pseudotsuga
menziesii 189.8 619 39.3 100 3.100 469 162 110 157 100 64 21 5 43.3
Tsuga
heterophylla 63.5 154 8.7 62 6,600 429 38 40 33 21 17 5 0 35.1
Thuja
plicata 46.7 123 6.6 43 900 374 50 19 38 2 7 5) 2 442
Total
all species 300.0 896 54.6 — 10,600 1272 250 169 228 123 88 31 7 —
*Importance value = Relative density + relative basal area + relative frequency (after Curtis and McIntosh 1951: Bray and Curtis 1957: Curtis 1959).
Figure 3. Eurhynchium oreganum completely
dominates the bryophyte layer with a mean
coverage of 23%. Hylocomium splendens is the
second most important species with a mean
coverage of 5%.
Discussion
Pseudotsuga menziesii is the dominant tree
species, constituting the uppermost layer of the
forest canopy. This species is suitably adapted to
all possible moisture and nutrient levels con-
trolled by the soil and topography on this
reserve. The highest productivity of the species
Tree and High Shrub Stratum
Pseudotsuga menziesii
Isuga_ heterophylla
Thuja olicata
Low Shrub Stratum
Gaultheria, shalion
Berberis nervosa
Rosa gymnocarpa
High Herb Stratum
Pteridium, aquilinum.
Polystichum munitum
in this dry subzone of the CDF zone is a site
index of 45 to 48 mat 100 yr. Trees near the base
of the slope on this reserve will probably attain
this size class at 100 yr. Because this dry subzone
is the least productive forest area of all the
coastal parts of the mesothermal climatic region
in British Columbia (Krajina 1969), Pseudot-
suga menziesii does not achieve the growth
reached by this species in more favorable
regions.
Tsuga heterophylla (Western Hemlock) is in
low abundance and, in general. trees of this
species are smaller than those of Pseudotsuga
30 40 . +50... GCiaae
Average percent cover
FIGURE 2. Average percent cover values for tree, low-shrub, and high-herb strata in the Saturna Island Ecological Reserve,
based on estimates in plots and subplots.
SS)
Low Herb-Dwarf Shrub Stratum
Festuca occidentalis
Festuca sSubulata’
Galium aparine
Elymus glaucus
Trientalis latifolia
Fragaria vesca
Listera cordata
Bryophyte Stratum
Eurhynchium oreganum
Hylocomium splendens
Dicranum fuscescens
Scapania nemorgsa
Isothecium _stoloniferum
Rhytidiadelphus triquetrus
Hypnum circinale
Lepidgzia reptans
Rhizomnium glabrescens
Lophocolea bidentata
O
NO
SULLIVAN: DOUGLAS FIR FOREST, B.C.
129
aS
6
Average percent cover
FIGURE 3. Average percent cover values for low-herb — dwarf-shrub and bryophyte strata in the Saturna Island Ecological
Reserve, based on estimates in subplots.
menziesii. Most Western Hemlock seedlings and
saplings are growing on rotting wood compared
with the soil or litter habitat characteristic of
Douglas Fir seedlings. This low density and
growth on decaying wood is typical of Tsuga
heterophylla in the dry subzone of the CDF
zone, which generally presents limiting factors
for the growth of this species. It has a very high
shade tolerance and prefers soils which supply
low amounts of fairly well balanced nutrients,
and these conditions exist only on podzolized
soils frequently moistened by rainfall (Krajina
1969). Adequate precipitation (at least 165 cm
annually) and a true podzol soil are not present
on the ecological reserve, thereby limiting the
success of Tsuga heterophylla.
Thuja plicata is found downslope in the
northern half of the reserve in association with
Tsuga heterophylla and the dominant Pseudot-
suga menziesii. It also occurs sporadically in
clumps in local topographic depressions where
some surface water is present in all seasons
except summer. This species has smaller trees ©
than Pseudotsuga menziesii but does grow very
well at lower elevations. The shade tolerance of
Thuja plicata is as high as that of Tsuga
heterophylla but its other ecological charac-
teristics are quite different (Krajina 1969).
Precipitation and soil texture, as they affect
availability of water and nutrients, are probably
limiting factors for Thuja plicata and Tsuga
heterophylla in the Douglas Fir forest com-
prising this ecological reserve. Pseudotsuga
menziesii 1s well adapted to subhumid or even
dry climates and if its moderate nutritional
requirements are met, it tends to be very
successful in both dominance and abundance,
relative to other species in this study area.
130
The associations of the Coastal Douglas Fir
zone have been studied and described by
Mueller-Dombois (1959) with reference to the
Nanaimo Lakes region and adjacent valleys. But
there have been no plant community studies on
this biogeoclimatic zone on the Gulf Islands.
Three of the seven recognized associations of
the Coastal Douglas Fir zone (dry subzone) are
present on this ecological reserve. The Salal,
moss, and Sword Fern (Polystichum munitum)
associations are all represented. The Salal
association (ca. 65% relative area) is most
widespread and occurs with all three tree species.
Herbs and bryophytes are suppressed owing to
the presence of Gaultheria shallon. The moss
association (ca. 25% relative area) alternates
with Salal in a patchy distribution throughout
the reserve. Eurhynchium oreganum and Hylo-
comium splendens best represent this com-
munity in their appearance as mats on the forest
floor.
The Sword Fern association (ca. 10% relative
area) is probably best represented on this reserve
at lower elevations. Here Pseudotsuga menziesii
achieves its best growth in both height and
diameter. Thuja plicata also follows this trend
but is still the subdominant species. Polystichum
munitum is not abundant in the forest but
exhibits its highest cover value in this part of the
reserve.
A brief discussion of the probable history and
future successional status of this forest is of
importance to the understanding and manage-
ment of this ecological reserve. There are a few
isolated individuals of Pseudotsuga menziesii
scattered throughout the forest, which are much
larger and older than the majority of trees. Most
of these trees, dead and alive, show evidence of
fire damage. Also, traces of burnt logs and wood
are distributed over the reserve. Therefore, at
least part of the area has experienced past fire,
which probably affected the growth of trees in
the present-day forest. It is possible that the very
large trees that survived the fire provided a seed
source for regrowth of the Douglas Fir forest.
Logging in the past is evident on the extreme
eastern part of the reserve but is definitely absent
from all other areas.
The future forest will continue to be dom-
inated by Pseudotsuga menziesii. Its seedlings
are the most evenly distributed of all species and
THE CANADIAN FIELD-NATURALIST
Vol. 93
exhibit shade tolerance on all available habitats.
The high density of seedlings and saplings of
Tsuga heterophylla and Thuja plicata is due to
clumping in restricted habitats, which contrasts
with the widespread occurrence of Pseudotsuga
menziesii. Both these species will remain
restricted to specific sites in correlation with
water and nutrient requirements.
In the understory, the pattern of Salal,
mosses, and open areas should continue to
predominate, with suppressed herb and reduced
shrub layers due to the shade cast by Gaultheria
shallon. It is possible that Salal may expand and
shade out regions of the moss association if soil
moisture and topographic variation do not
continue to support this Salal-moss distribution.
Finally, the browsing of vegetation by
Columbian Black-tailed Deer (Odocoileus
hemionus columbianus) may become important
if the population density of these animals
increases. My sightings of deer during fieldwork
indicated that these animals were very abundant
throughout the reserve and on Saturna Island in
general. The understory species, Gaultheria
shallon, Rosa gymnocarpa, Pteridium aqui-
lium, and = Polystichum munitum exhibited
some degree of damage from browsing. Effects
on tree seedlings were not noticeable, but
according to J. Revel (1963 unpublished report,
Faculty of Forestry, University of British
Columbia) and Cowan and Guiget (1970), this
species of deer prefers seedlings of Western Red
Cedar and Douglas Fir. Western Hemlock is
usually exempt from browsing. Therefore, the
future successional status of this forest could be
controlled by the effects of seedling browsing as
well as the ecological tolerances already dis-
cussed for the three tree species. In addition, deer
trails on some downslope regions of the reserve
have disturbed the ground cover of mosses
exposed to the passage of these animals.
Acknowledgments
I thank Charles J. Krebs for reviewing the
manuscript and Charles E. Beil and the Ecolo-
gical Reserves Committee through V. J. Krajina
for financial support. Many thanks are extended
to Walter R. Kaiser and family for help with the
fieldwork. I am most grateful to Nancy Turner
and William Zales for identification of vascular
plants and bryophytes.
Ig
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Curtis, J. T. and R. P. McIntosh. 1951. An upland forest
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1: 1-17.
Krajina, V.J. 1969. Ecology of forest trees in British
Columbia. Ecology of Western North America 2: 1-147.
Krajina, V. J. 1973. The conservation of natural ecosystems
in British Columbia. Syesis 6: 17-31.
La Roi, G. H., T. A. Babb, and B. J. Perley. 1976. Cana-
dian National Directory of IBP areas. University of
Alberta Press, Edmonton.
Lawton, E. 1971. Moss flora of the Pacific Northwest.
Hattori Botanical Laboratory. Nichinan, Miyazaki,
Japan.
Mueller-Dombois, D. 1959. The Douglas Fir associations
on Vancouver Island in their initial stages of secondary
succession. Volume, I, 2. Ph. D. thesis, University of
British Columbia, Vancouver.
Stotler, R. and B. Crandall-Stotler. 1977. A checklist of
liverworts and hornworts of North America. Bryologist
80: 405-428.
Received 27 June 1978
Accepted 2 November 1978
Colonial-nesting Herring Gulls and Common Terns
in northeastern Saskatchewan
HARRY A. STELFOX! and GREGG J. BREWSTER?
'Wildlife Research Division, Saskatchewan Department of Tourism and Renewable Resources, 2602 8th Street East,
Saskatoon, Saskatchewan S7H 0V7
2Habitat Protection and Development Division, Saskatchewan Department of Tourism and Renewable Resources, 2602 8th
Street East, Saskatoon, Saskatchewan S7H 0V7
Stelfox, Harry A. and Gregg J. Brewster. 1979. Colonial-nesting Herring Gulls and Common Terns in northeastern
Saskatchewan. Canadian Field-Naturalist 93(2): 132-138.
Observations of colonial-nesting birds were conducted during the summers of 1973 and 1974 as part of the Wildlife
investigations of the Churchill River Study in northeastern Saskatchewan. Herring Gulls (Larus argentatus) and Common
Terns (Sterna hirundo) were the only species found to breed in abundance within the surveyed area. All of the Herring Gull
colonies and the majority of the Common Tern colonies were located on Reindeer Lake, rather than on the smaller lakes and
river channels along the Churchill and Reindeer rivers. The mean size of nesting colonies was relatively small.compared to
those of more southerly latitudes. Favored nesting sites were small rocky islands which were very susceptible to flooding.
Common Terns appeared to be better adapted to utilizing the more ephemeral nesting habitat along the river systems than
Herring Gulls. ~
Key Words: Common Terns, Herring Gulls, nesting colonies, habitat, behavior, Saskatchewan.
The breeding range of the Common Tern
(Sterna hirundo) extends throughout Saskat-
chewan and that of the Herring Gull (Larus
argentatus) covers most of the northern half of
the province (Gollop 1969). This paper presents
gull and tern nesting observations made during
the summers of 1973 and 1974 while we were
conducting wildlife studies in northeastern
Saskatchewan for the Churchill River Study
(CRS). The CRS was undertaken by the
governments of Canada, Manitoba, and Saskat-
chewan to assess the potential environmental
impact of a proposed hydroelectric dam on the
Churchill River 65 km west of the Manitoba-
Saskatchewan border (Barber et al. 1975).
The study area lies 300 km northeast of Prince
Albert, Saskatchewan, and is characterized by
Precambrian rock and boreal forest. The area
investigated includes the Churchill River from
Drinking Falls to the Island Falls Dam,
Manawan Lake, all of the Reindeer River and
most of Reindeer Lake (Figure 1).
The primary objective of our gull and tern
investigations was the location of nesting
colonies to determine population levels and
distribution. A secondary objective was an
assessment of the nesting habitat.
Methods
High-level black-and-white air photos were
examined to locate small isolated islands and
shoals representing potential nesting sites. In
late June 1973 an aerial survey was flown over
most of Reindeer Lake to check these locations
and numerous others for signs of nesting
activity. Nest sites on the Churchill and Reindeer
rivers were located during the course of other
wildlife aerial and ground surveys.
Once during late June or early July of each
year a sample of nesting islands was examined
on the ground to determine numbers of nests,
eggs, young, and adults, as well as nesting
habitat characteristics. Colonies on Reindeer
Lake were examined between 19 and 23 June in
both years. In 1974 elevations of islands and
individual nests above water level were surveyed
at 10 selected colony sites using a Cooke survey
transit and stadia rod. Nest distribution and
substrate characteristics were also noted.
Results
Common Terns
Twenty-seven colonial nesting sites were
located, 17 on Reindeer Lake, 4 on the Churchill
River, 4 on the Reindeer River, and 2 on
132
1979 STELFOX AND BREWSTER: COLONIAL BIRDS, SASKATCHEWAN 133
@ Common Tern Nesting Colony
A Herring Gull Nesting Colony
® Common Tern and Herring Gull Nesting
Colonies on the same island
Water bodies surveyed for colonial
nesting birds
wwe ose
Ug h\s
3
SASKATCHEWAN
Prince
albert
OSaskatoon
ORegina
VEOLINVW
6 Southend
Horriot R.
Ww?
NMUMIHILVASYS
FIGURE 1. Location of Common Tern and Herring Gull nesting colonies within the study area.
134
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE |—Number and average size of Common Tern and Herring Gull nesting colonies
Number of colonies
located (and sampled)
Mean number (and range) of Mean number (and range) of
adults per sampled colony nests per sampled colony
Location 1973 1974 1973 1974 1973 1974
Common Terns
Reindeer Lake 13(5) 15(6)* 23(10-45) 25(6—5S0) 12(4-26) 21(5-38)
Reindeer River 2(2) 3(3) =" 34(18-—50) 55(35-66) 19(9-28) 25(11-33)
Churchill River 4(3) flooded 28(8-45) = 14(9-22) =
Manawan Lake 2(2) not surveyed 43(40-45) — 29(17-40) —
Means 29 35 16 23
Herring Gulls
Reindeer Lake 9(5) NG) 19(10-S5) 18(6—45) 13(2-31) 13(3-22)
*Includes five colonies not sampled in 1973.
**Includes two colonies not sampled in 1973.
***Includes four colonies not sampled in 1973.
Manawan Lake (Figure 1). Not all colony sites
were used in both years. In 1974, colony sites on
the Churchill River were completely flooded asa
result of unseasonally high water levels.
Mean colony size was small, ranging from 16
nests in 1973 to 23 in 1974 (Table 1). The two
colonies on Manawan Lake were relatively
large, averaging 29 nests in 1973. Colonies
examined on Reindeer Lake averaged only 12
nests in 1973 and 21 nests in 1974.
By 21 June, most colonies had nests with eggs,
with mean clutch sizes ranging from | to 2.8. On
20 June 1973, however, two colonies on Rein-
deer Lake had several adult birds and nest
depressions with no eggs, whereas a colony on
the Reindeer River had the first young hatching
on 23 June 1973. Reliable productivity data were
not obtained because most sampled colonies
were visited only once during the season.
Some loss of eggs from flooding was observed,
with low near-shoreline nests being washed over
during storms and strong winds. Heavy rains
which resulted in sharply rising water levels also
took their toll. One colony on Reindeer Lake
examined on 23 June 1974 had recently been
destroyed by humans: 10 adult terns remained
on the shoal but no re-nesting had been initiated.
Herring Gulls
Eleven different colony sites were located, all
on Reindeer Lake. Four of these were shared
with Common Terns (Figure 1). In addition, a
single Herring Gull nest with eggs was at a
Common Tern colony of Steephill Lake (Rein-
deer River) in both 1973 and 1974, and lone
Herring Gull nests were observed at four
locations on Reindeer Lake, three of these being
associated with Common Tern nesting sites.
Colonies averaged 13 nests in 1973 and 1974
(Table 1). In 1973 most nests contained eggs,
while in 1974 a large percentage of the nests
attended by adults contained no eggs.
Nest Island Characteristics
All Herring Gull and Common Tern nesting
islands were very low (<3 m above water level)
and small (<1000 m’), except for one larger
Herring Gull nesting island (Colony #10, see
Table 2). Generally less than 50% of the surface
of nest islands supported vegetation, primarily
grasses, forbs, mosses and lichens, with an
occasional shrub or tree. But Colony #10 in
Table 2 and a Common Tern colony on the
Reindeer River (Figure 2) had 70-80% of their
surface area covered by shrubs and trees,
whereas several islands had no growing vegeta-
tion (Figure 3). The vegetation and small
amount of associated soil occurred on the higher
portions of nesting islands where there was no
extensive erosion by ice and wave action.
The substrate of nesting islands varied from
large fragmented blocks to smoother solid rock
and scattered boulders and stones. Occasionally
some finer sand and pebble deposits were
present near the water’s edge. Driftwood was
strewn over the lower portions of some islands
and completely covered one (Colony #4, Table
)).
~ 1979 STELFOX AND BREWSTER: COLONIAL BIRDS, SASKATCHEWAN 135
FIGURE 2. Common Tern nesting colony on Steephill Lake (Reindeer River).
EMSs
SN Ds
a
\
ee SS
FiGuRE 3. Herring Gull colony.
136
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 2—Island size and nest distribution for 10 Herring Gull and Common Tern nesting islands on Reindeer Lake, 1974
Nest height above water (m)**
Island Island
Colony height (m) area (m2) Species
l 1.6 155 Common Tern
2 Dl 349 Common Tern
Herring Gull
3 1.0 697 Common Tern
Herring Gull
4 0.8 387 Common Tern
Herring Gull
5 3.1 960 Common Tern
Herring Gull
6 2.4 968 Common Tern
Herring Gull
7 2.4 465 Herring Gull
8 2S) 426 Herring Gull
9 Dal 426 Herring Gull
10 2.0 3988 Herring Gull
Means 1.8 586 Common Tern
Dal 1083 Herring Gull
Number
of nests* Mean Range
5) 0.4 0.3-0.4
19 0.6 0.4-0.8
l 1.3
38 0.4 0.2-0.5
| 1.0
16 0.2 0.1-0.4
3 0.5 0.2-0.8
36 1.2 0.2-2.0
I 0.2
14 0.9 0.8-1.1
2D 0.8 0.5-1.1
22 0.5 0.1-0.8
6 0.5 0.3-0.8
10 0.8 0.5-1.4
13 0.5 0.2-1.1
0.7
0.7
*All recognizable nests, with or without eggs, were included in the measurements.
** All measurements were taken from 21 to 23 June 1974 when the water level on Reindeer Lake was 331.1 m (1103.8 ft.) above sea-level.
Distribution of Nest Sites
Common Tern nests were usually located near
the lower edge of the vegetated portions of
islands among scattered grass tussocks and
moss-lichen mats. Nests were sometimes nothing
more than a shallow depression in a thin layer of
soil, or a crevice in the rock. Herring Gulls
constructed nests of grasses and small twigs,
frequently on exposed rocks or pieces of
driftwood, affording a good vantage point.
The mean height of both tern and gull nests
above late June water levels in 1974 was 0.7 m
(Table 2). No nests were located more than
2.0 m above water level. The mean nearest-
neighbor distance between the edges of
Common Tern nests was 0.9 m, with the closest
nests being only 0.3 m apart. The mean distance
between adjacent Herring Gull nests was 3.3 m,
while the shortest distance was 2.1 m.
Discussion and Conclusions
Herring Gulls and Common Terns were the
only species of Laridae found to breed in
abundance within the study area. All of the
Herring Gull colonies and most of the Common
Terns were found on Reindeer Lake rather than
the smaller lakes and river channels to the south
(cf., Vermeer 1975). This distribution is thought
to reflect the more stable nesting conditions on
Reindeer Lake, where water levels are not
subject to rapid large-scale fluctuations such as
those that occur on the rivers.
Common Tern colony locations and sizes on
the Churchill and Reindeer rivers changed from
1973 to 1974 because of late spring flooding in
1974. McNicholl (1975) points out that in
unstable habitats larids generally demonstrate
reduced nest site tenacity and increased group
adherence, thus aiding the rapid pioneering of
alternate nesting habitat. The shifting nest site
locations and relatively larger average size of
Common Tern colonies that we observed in 1974
on the river system support this theory. We
observed no evidence of Common Terns nesting
singly, while we found five such cases in Herring
Gulls. Several other cases of Herring Gulls
nesting as single pairs have been reported in the
literature (Harper 1953; Nero 1963; Vermeer
1975). In general our data would suggest that
nest site tenacity is stronger in the Herring Gulls
while group adherence is stronger in the
Common Terns, thus enabling the terns more
successfully to colonize riverine habitats.
A noteworthy feature of our tern and gull
nesting colonies is their small mean size. This
appears to be characteristic for these more
1979
northern latitudes as indicated by other workers
(Mowat and Lawrie 1955; Scotter 1961; Nero
1963, 1967: Weber 1976). No colonies of over
100 nests were reported for either species and
most were considerably smaller. By contrast,
Common Tern colonies in southern Saskat-
chewan are characteristically large, ranging in
size from 100 to 1000 nests (Ferry 1910;
Lahrman 1957; Sanderson 1966; C. S. Houston,
personal communication). In the Great Lakes,
Common Tern colonies may number over 2500
birds, and Herring Gull colonies usually exceed
100 birds (Morris and Hunter 1976). Vermeer
(1975) studied one Herring Gull colony of 161
nests at Kawinaw Lake in southern Manitoba.
Several factors may account for such variation.
Differential biological productivity and food
availability is likely a primary factor (cf.,
Vermeer 1970). The waters in our study area are
relatively nutrient-poor (oligotrophic) and have
a cover of ice until mid- to late May. The size
and distribution of nest islands, plus predation
factors, may be important as well.
Favored nesting sites of both gulls and terns
were very low, small, rocky islands with
restricted vegetation growth. Wind, wave and ice
action are believed to be important elements in
maintaining the exposed mineral substrates and
sparse vegetation cover utilized for nesting. Nest
sites thus occurred within a few feet of sur-
rounding water levels and were very susceptible
to flooding from both natural and artificial
water level increases.
A hydroelectric dam and reservoir as pro-
posed by Saskatchewan Power Corporation at
Wintego Rapids* on the Churchill River would
result in water levels rising 28.5 m at Wintego
Lake, 5.3 m at Steephill Lake, and 2.3 mat Keg
Lake. All eight of the islands used for nesting by
Common Terns on the Churchill and Reindeer
rivers would be flooded; however, at least 11 new
potential nesting islands would be created by the
flooding. Reindeer Lake, which would be in-
cluded within the proposed reservoir, would not
exceed its historical levels, resulting inno change
in numbers and locations of nesting islands.
Once filled and operational, reservoir water
levels would steadily rise a full 1.2 m from 1 May
to | November to replace the winter drawdown.
Much of this rise would come in June and July
when runoff is heaviest (Hofer 1975). Our data
STELFOX AND BREWSTER: COLONIAL BIRDS, SASKATCHEWAN 137)
on the distribution of gull and tern nests (Table
2) suggests that on suitable nesting islands within
the proposed Wintego Reservoir a 0).7-m in-
crease in water levels during June and July
would flood more than 50% of the gull and tern
nests.
Further studies are needed at these northern
latitudes to understand more fully the factors
influencing population distributions and pro-
ductivity. It is necessary to have a better
understanding of behavioral responses to
changing environmental conditions, as well as
food and nesting habitat requirements, before
one can reasonably predict the possible impacts
of future northern developments on colonial-
nesting gulls and terns.
Acknowledgments
This study was funded by, and carried out for,
the Churchill River Study under the direction of
the Wildlife Research Division, Department of
Tourism and Renewable Resources, Govern-
ment of Saskatchewan. Valuable field assistance
was provided by D. G. Hjertaas, D. G. McInnes,
Re Wo Melinchuk: J. Es Polson. vand | Dsair
Sherratt. We thank C.S. Houston for many
helpful suggestions and critical review of the
manuscript.
*On 18 September 1978 the Government of Saskatchewan,
following the recommendation of the Churchill River
Board of Inquiry, announced that it would not permit
construction of the proposed hydroelectric installation
at Wintego Rapids.
Literature Cited
Barber, S. R., H. A. Stelfox, and J. D. Boden. 1975. Wild-
life (Saskatchewan). Churchill River Study, Final
Report 28. Saskatoon, Saskatchewan. 272 pp.
Ferry, J. F. 1910. Birds observed in Saskatchewan during
the summer of 1909. Auk 27: 185-204.
Gollop, J.B. 1969. Birds in Saskatchewan. Jn Atlas of
Saskatchewan. Edited by J.H. Richards and K. I.
Fung. Modern Press, Saskatoon. pp. 85-90.
Harper, F. 1953. Birds of Nueltin Lake Expedition. Ameri-
can Midland Naturalist 49: 1-116.
Hofer, R. D. 1975. Hydrology (Saskatchewan). Churchill
River Study, Final Report 2. Saskatoon, Saskat-
chewan. 61 pp.
Lahrman, F. W. 1957. Birds of the Isle of Bays. Blue Jay
15: 106-110.
McNicholl, M.K. 1975. Larid site tenacity and group
adherence in relation to habitat. Auk 92: 98-104.
138
Morris, R. D. and R.A. Hunter. 1976. Factors influenc-
ing desertion of colony sites by Common Terns (Sterna
hirundo). Canadian Field-Naturalist 90: 137-143.
Mowat, F. M. and A. H. Lawrie. 1955. Bird observations
from southern Keewatin and the interior of northern
Manitoba. Canadian Field-Naturalist 69: 93-116.
Nero, R. W. 1963. Birds of the Lake Athabasca region,
Saskatchewan. Saskatchewan Natural History Society,
Special Publication Number 5. 143 pp.
Nero, R.W. 1967. The birds of northeastern Saskat-
chewan. Saskatchewan Natural History Society,
Special Publication Number 6. 96 pp.
Sanderson, R. M. 1966. The colonial birds at Suggi Lake,
Saskatchewan in 1966. Blue Jay 24: 121-123.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Scotter, G. W. 1961. Summer observations of birds in
northern Saskatchewan, 1960. Blue Jay 19: 70-74.
Vermeer, K. 1970. Breeding biology of California and
Ring-billed Gulls. Canadian Wildlife Service, Report
Series Number 12. 37 pp.
Vermeer, K. 1975. Food habits and breeding range of
Herring Gulls in the Canadian prairie provinces.
Condor 75: 478-480.
Weber, W. C. 1976. Birds of north-central Manitoba, 1973.
Blue Jay 34: 84-94.
Received 10 August 1978
Accepted 23 November 1978
Epiphytes on White Elm, U/mus americana,
near Thunder Bay, Ontario
P. BARCLAY-ESTRUP! and RICHARD A. SIMS?
'Department of Biology, Lakehead University, Thunder Bay, Ontario P7B 5El
2Canadian Forestry Service, Great Lakes Forest Research Centre, Sault Ste. Marie, Ontario P6A 5M7
Barclay-Estrup, P. and R. A. Sims. 1978. Epiphytes on White Elm, U/mus americana, near Thunder Bay, Ontario.
Canadian Field-Naturalist 93(2): 139-143.
White Elm (U/mus americana) stands near Thunder Bay, Ontario were investigated for epiphyte cover. Lichens dominated
the cover in upper quadrats and mosses dominated in quadrats near the ground. In both upper and lower quadrats, mean
cover was about 10%. Total cover was greatest on the north sides of the trees and tended to increase as the distance from the
city increased. The stand located close to a kraft pulp and paper mill had the lowest number of species. Three species of
liverworts, 12 mosses, and 17 lichens were recorded.
Key Words: northern Ontario, White Elm, epiphytes, pollution.
White Elm, U/mus americana, is rapidly
disappearing from much of its former range in
Canada and the United States owing to large-
scale mortality caused by the introduced Dutch
Elm Disease. This unfortunate circumstance has
resulted in increased interest in studies relating
to White Elm, such as that of Mahoney (1973) 1n
northern Michigan and Newberry (1974) in
Wisconsin. A further point of interest is that the
groves in our region, the northwest shore of
Lake Superior, form part of the northern
boundary of White Elm’s range. The purpose of
this study was to record the epiphytes on White
Elm near Thunder Bay before Dutch Elm
Disease removes the elm population, and to note
epiphyte variability in relation to distance from
the city of Thunder Bay.
Methods
During September and October of 1973, 50
trees were examined in five natural stands (10
trees per stand) located at various distances from
the city of Thunder Bay. These stands were as
follows:
1. Great Lakes Paper. On the south bank of the
Kaministiquia River directly south of the
Great Lakes Paper kraft pulp and paper mill.
This small stand is on the south end of
Thunder Bay city and is considered to be at
0 km (48°21’W, 89°19’N, 194 m above sea-
level (asl)).
2. Pointe de Meuron. This relatively large stand
of elm is on the north bank of the Kaminis-
tiquia River adjacent to the reconstructed
Old Fort William and is 3 km west of the
Great Lakes Paper site (48°21’W, 89°21’/N,
195 m asl).
3. Slate River. This medium-sized stand is on
the west bank of the Slate River just north
of Highway 608 and is 16 km southwest of
Great Lakes Paper (48°16’W, 89°29’N,
224 m asl).
4. Kakabeka Gorge. This is a small stand
located on the east bank of the Kaminis-
tiquia River about 300 m downriver from
Kakabeka Falls (36 m high) and is 23.4 km
west-north-west of Great Lakes Paper
(48°24’W, 89°37'N, 229 m asl).
5. Blind Line Road. This small stand is on the
south bank of a small creek just north of
Blind Line Road and 250 m west of High-
way 11-17. This site is 25.6 km west-north-
west of Great Lakes Paper (48°25’W,
89°38’N, 284 m asl).
All sites have Balsam Poplar (Populus bal-
samifera), Black Ash (Fraxinus nigra), Moun-
tain Maple (Acer spicatum), and Red-osier
Dogwood (Cornus stolonifera). Also generally
present are Eastern White Cedar (Thuja occi- ——
dentalis), White Birch (Betula papyrifera),
Speckled Alder (A/nus rogosa), and Red Rasp-
berry (Rubus idaeus). Table | shows additional
characteristics of the sites and of the trees in the
stands.
139
140 THE CANADIAN FIELD-NATURALIST Vol. 93
TABLE 1—Mean soil and tree characteristics. DBH = Diameter at breast height (range in parentheses)
Site Soil pH % Carbon Trees, DBH (cm) Bark pH
Great Lakes Paper 5.9 11.0 44.9 (30.5-71.1) 4.2
Pointe de Meuron 5.9 13.1 37.7 (19.7-55.9) 6.2
Slate River 5.8 14.4 38.0 (22.9-49.5) 6.4
Kakabeka Gorge 6.8 5.3 24.8 (17.2-45.7) 6.5
Blind Line Road 6.4 18.8 21.3 (15.9-31.8) 6.6
Mean 6.2 IDES 381.3} OS 9=7 11 1b) 6.0
TABLE 2—Presence (X) and frequency of epiphytes on White Elm at the five study sites. Frequency figures are out of a
possible total of 50 trees. ¢ = found only in top quadrats: b = found only in bottom quadrats: no notation = found in both top
and bottom quadrats. GLP = Great Lakes Paper: PM = Pointe de Meuron: SR = Slate River: KG = Kakabeka Gorge:
BL = Blind Line Road
Presence Frequency
Top Bottom Whole
Species GLP PM SR KG BL quadrats quadrats tree
Frullania eboracensis Xx Xx xX xX 28 15 31
Frullania bolanderi x 5 | 5
Porella platyphylla xe xe 0 2 D
Anomodon minor x? x xX x 6 20 20
Orthotrichum speciosum x x! x 15 Fi 19
Brachythecium salebrosum x x xX 8 13 15
Pylaisiella selwynii Xx X X 5 7 1]
Pylaisiella polvantha x! x 10 10 11
Brachythecium reflexum x xe x 3 3 3
Platydicta subtile xX | 5 5
Plagiothecium denticulatum Xe, 0) 2 2
Amblystegium juratzkanum x? 0 l l
Orthotrichum obtusifolium x! | 0 l
Dicranum montanum x 0 I |
Mnium cuspidatum xe 0 I l
Physconia grisea x xX xX x xX 30 23 33
Candelaria concolor xe Xx xX x! 28 10 29
Parmelia sulcata xX xX x! xX 21 9 21
Physcia orbicularis xX xX xX x 18 6 19
Physcia adscendens x x! x’ 11 3 12
Physcia aipolia xX x! Xx 7 3 9
Bacidia chlorococca* X 4 7 8
Lepraria membranacea xX 3 4 5
Lecanora impudens x X xX 3 3 4
Lobaria pulmonaria Xx x! Xx 4 l 4
Xanthoria polycarpa x! x! 4 0 4
Cetraria ciliaris x! 3 0 3
Lecanora conizaea xX 2 2 3
Evernia mesomorpha x! 2 0 2
Lepraria neglecta X | l 2
Parmelia saxatilis x! 2 0 2
No. of species, top 4 18 [2 12 ae)
No. of species, bottom 6 16 9 10 13 XG=aMil
No. of species per tree 6 20 12 15 IG SNe
* Bacidia chlorococca, while not noted in the quadrats. was also present on the elms at both Slate River and Blind Line Road.
1979
Each of the 10 trees ineach stand was analyzed
for epiphyte presence using the method em-
ployed by Rao and Le Blanc (1967). In this
method, two circular quadrats were used per
tree. One quadrat (bottom) was from the tree
base at 0 cm to 30 cm above the base; the second
quadrat (top) was also 30cm deep and was
located at 120 cm to 150 cm above the tree base.
All mosses, liverworts, and lichens were
recorded. Cover was also estimated using the
Domin Scale for this purpose. Conversion of the
Domin Scale data to percent cover was based on
the following equivalents: 10 = 100%, 9 = 75%,
8 = 60%, 7=40%, 6= 30%, 5 = 20%, 4=5%,
3=3%, 2=1%, 1=0.5% and K =0.25%. The
directional quarter of greatest cover for each tree
was noted.
Sorensen’s Coefficient of Similarity was used
to compare quadrats and trees and also as a basis
for a Bray and Curtis (1957) ordination in which
each tree (top and bottom combined) was used
as one quadrat.
All field studies were done by the second
author. Determinations were also done by the
second author with assistance from C. Garton
(bryophytes) and the first author (macro-
lichens). All crustose lichens were verified by
J.W. Thompson. Voucher specimens were
SITES
Great Lakes
Point de Meuron
Slate River
Kakabeka Falls
Blind Line Road
| @
Uy ZO
Wi@ 37% 3 A
‘ 4a
5 #
FIGURE |. An ordination of whole trees from five sites using
the Sorensen Coefficient of Similarity method of
Bray and Curtis (1957).
BARCLAY-ESTRUP AND SIMS: WHITE ELM EPIPHYTES 14]
deposited in the Lakehead University Her-
barium.
Results
The species of lichens and bryophytes present
at each site, the number of species per tree, the
number of species in top quadrats, the number of
species in bottom quadrats, and the number of
species per site in all quadrats are shown in Table
2. The total number of species for all sites is 32,
made up of 3 hepatics, 12 mosses, and 17
lichens.
The coefficient of similarity ordination of
whole trees is illustrated in Figure | to show
relationships between sites.
Percent cover values by site are shown in
Figure 2. When all five sites are considered
40
30
20
% COVER
FIGURE 2. Cover relationships of bryophytes and lichens
showing values for top quadrats (above line) and
bottom quadrats (below line) at the five study sites.
The sites are arranged in order of increasing dis-
tance from Thunder Bay. L, lichens: B, bryophytes:
T, total of both lichens and bryophytes: GLP,
Great Lakes Paper site: PM, Pointe de Meuron site:
SR, Slate River site: KG, Kakabeka Gorge site:
BR, Blind Line Road site.
142
together, lichens contribute a mean cover of
5.9% in the top quadrats and 2.1% inthe bottom
quadrats. Byrophytes contribute 3.6% in the top
quadrats and 8.0% in the bottom quadrats.
Overall mean epiphyte cover was 9.5% in the top
quadrats and 10.2% in the bottom quadrats. In
68% of the trees the greatest cover 1s on the north
quarter of the trees, 14% on the west, 10% on the
east, 2% on the south, and in 6% two or more
quarters were equal.
Discussion
The most obvious observation from the data
obtained in this study is that the Great Lakes
Paper site is quite different from the other four
areas. The number of species of epiphytes in each
of the three groups, and the total number of
species for all groups, are significantly lower
(P< 0.01) in the Great Lakes Paper site than in
any of the other areas. This is perhaps even more
significant when related to the mean DBH
(diameter at breast height) of the stands, as
Great Lakes Paper has the highest mean DBH of
any of the areas, indicating greater age which
often results in high epiphytic cover. The bark
pH of 4.2 of the Great Lakes Paper site was
much lower than on the other sites, which had a
mean bark pH of 6.4. The emissions from the
nearby pulp mill are probably responsible for the
singular character of this site when compared to
the other four sites. The mean bark pH of 6.0 is,
incidentally, comparable to that found by Skye
(1968) for U/mus sp. in the Stockholm region,
and also for White Elm in Central Wisconsin
(Newberry 1974).
When all sites are considered together as
representative of the Thunder Bay region, the
number of species of lichens compares favorably
with the number noted by Mahoney (1973) on
White Elm in northern Michigan, where 15
species were found in one area and 8 species in
another for a total of 17.
Other publications that have included data on
White Elm epiphytes are Hale (1952, 1955),
Culberson (1955), Le Blane and De Sloover
(1970), and Newberry (1974). For various
reasons (e.g., data for elm are combined with
that of other species: whole trees were investi-
gated: trees were of urban environments) com-
parison with all but Newberry (1974) are perhaps
not very useful, except for the obvious con-
THE CANADIAN FIELD-NATURALIST
Vol. 93
clusion that generally the more trees and the
larger the area of the tree included, the greater
the number of epiphyte species that will be
encountered. Newberry’s (1974) study, however,
does give separate White Elm data and is related
to sulphur dioxide emissions from a kraft paper
mill. In Newberry’s study a species list is not
included but 18 species and three varieties of
lichens are common on White Elm and eight
more lichen species are classified as rare (six
more fruticose lichens are present but the
phorophyte is not specified). Candelaria con-
color, Parmelia sulcata, Physcia orbicularis, s.1.,
and Physconia grisea are apparently common in
Thunder Bay as well as in northern Michigan
and central Wisconsin. Physcia setosa, the most
common lichen in the Michigan study, is rare in
Thunder Bay and was not noted on the elms
studied. Physcia elaeina, a ubiquitous species in
Wisconsin, has not been reported in the Thunder
Bay region. Parmelia rudecta and P. caperata
were present in both the American studies, but
although present in Thunder Bay, were not
recorded on the elms investigated. One of the
most common epiphytes in the Thunder Bay
area 1s Hypogymnia physodes. This species,
however, was not noted on the White Elms of
this study or by Mahoney (1973) in Michigan
and was reported as rare in Wisconsin, indicat-
ing if not phorophyte specificity at least phoro-
phyte preference.
Although the bryo-flora was not considered
by Mahoney (1973) or Newberry (1974) some
general comments are perhaps warranted. None
of the three liverworts found are rare. But, while
Frullania eboracensis, the most common epi-
phytic hepatic in this study, is also one of the
most common in the Thunder Bay region the
other two species are scarce. In the mosses,
moreover, Anomodon minor, the most common
moss on elm, is also scarce, and Hypnum
pallescens, usually a very common epiphytic
moss, was not recorded on any of the 50 elms
studied.
Table 2, besides showing the species present,
clearly indicates that liverworts and lichens are
more common in the upper quadrats and that
the mosses are more common in the lower
quadrats.
The ordination (Figure |) based on presence-
absence data also shows the Great Lakes Paper
1979
site to be rather different from the other four
sites. The Kakabeka Gorge site is set off as well,
but not to the same extent and the ordination
indicates relationships among the Pointe de
Meuron, Slate River, and Blind Line Road sites.
A second ordination using the reciprocal averag-
ing technique (Hill 1973) gave essentially the
same results except that the Kakabeka Gorge
stand is quite separate from the other groups.
The cover relationships between groups are
similar to the species numbers relationships,
with bryophytes being greater in the lower
quadrats and lichens greater in the upper
quadrats. Cover relationships between sites,
however, are quite different from species num-
bers relationships. In Figure 2 the sites are
arranged according to increasing distance from
Thunder Bay City. Coincidentally, this is in
decreasing order of tree DBH which probably
represents decreasing mean age of stands as well.
Increasing distance from Thunder Bay results in
increasing cover, attributable at least in part to
decreasing aerial poijlutants. The pattern is not
as straightforward as this, however, as the very
high bryophyte cover at Kakabeka Gorge must
be related to the higher humidity levels resulting
from spray from nearby Kakabeka Falls. In any
event the Great Lakes Paper site has the lowest
cover values and Blind Line Road, although it 1s
apparently the youngest stand, has cover values
equivalent to those of the larger-diameter trees
at both Pointe de Meuron and Slate River.
In conclusion it can be stated that the epiphyte
flora of White Elms near Thunder Bay 1s
relatively comparable to the floras of elms in
Michigan and Wisconsin. Also, if Dutch Elm
Disease, which was first reported near Thunder
Bay in 1976 (Anonymous 1976), should become
severe in these isolated stands of elms in this
northern part of the tree’s range, some record is
now available of the present epiphyte flora.
BARCLAY-ESTRUP AND SIMS: WHITE ELM EPIPHYTES 143
Acknowledgments
We thank W. M. Graham of the Lakehead
Biology Department for useful discussions:
J. W. Thompson, Department of Botany, Uni-
versity of Wisconsin for determining a number
of lichens: and the National Research Council of
Canada for support under Grant Number A-
6121.
Literature Cited
Anonymous. 1976. Tree diseases. Forestry Bulletin, June—
July. Great Lakes Forest Research Station, Sault Ste.
Marie, Ontario.
Bray, J. R. and J. T. Curtis. 1957. An ordination of the
upland forest communities of southern Wisconsin. Eco-
logical Monographs 27: 325-349.
Culberson, W. L. 1955. The corticolous communities of
lichens and bryophytes in the upland forests of northern
Wisconsin. Ecological Monographs 25: 215-231.
Hale, M.E., Jr. 1952. Vertical distribution of crypto-
gams in a virgin forest in Wisconsin. Ecology 33:
398-406.
Hale, M. E., Jr. 1955. Phytosociology of corticolous cryp-
togams in the upland forests of southern Wisconsin.
Ecology 36: 45-63.
Hill, M.O. 1973. Reciprocal averaging: an eigenvector
method of ordination. Journal of Ecology 61: 237-249.
Le Blanc, F. and J. De Sloover. 1970. Relation between
industrialization and the distribution and growth of
epiphytic lichens and mosses in Montreal. Canadian
Journal of Botany 48: 1485-1496.
Mahoney, R. 1973. The lichen flora on Ul/mus americana
in northern Michigan. Michigan Botanist 12: 163-166.
Newberry, G. 1974. The influence of a sulfate-process
paper mill on corticolous lichens. Bryologist 77: 561-576.
Rao, D.N. and F. Le Blanc. 1967. Influence of an iron-
sintering plant on corticolous epiphytes in Wawa, On-
tario. Bryologist 70: 141-157.
Skye, E. 1968. Lichens and air pollution, a study of
cryptogamic epiphytes and environment in the Stockholm
region. Acta Phytogeographica Suecica 52, Uppsala. '
123 pp.
Received 14 February 1975
Resubmitted 16 October 1978
Accepted 16 November 1978
Habitat Utilization and Population Densities of the
Amphibians of Northeastern Alberta
WAYNE ROBERTS and VICTOR LEWIN
University of Alberta Museum of Zoology, Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9
Roberts, W. and V. Lewin. 1979. Habitat utilization and population densities of the amphibians of northeastern Alberta.
Canadian Field-Naturalist 93(2): 144-154.
During 1976 and 1977 a survey of the herpetofauna of boreal forest habitats of northeastern Alberta was undertaken. The
Wood Frog (Rana sylvatica) was the most abundant and widespread species and was collected at 24 of 25 sites examined.
Boreal Chorus Frogs ( Pseudacris triseriata maculata) and Canadian Toads (Bufo hemiophrys) were common but found at
less than half of the study sites. All three species had spawned prior to 12 June. Natural ponds and borrow pits were the most
frequently used spawning sites. Canadian Toads also spawned in flowing water and lake margins. All species had
metamorphosed by early August. Wood Frogs metamorphosed at a mean snout—vent length of 17.2 mm, Canadian Toads at
12.4 mm, and Boreal Chorus Frogs at 13.7 mm. Spawning for each of these species probably does not occur until individuals
are in their third summer of life. Population densities of Wood Frogs, Boreal Chorus Frogs, and Canadian Toads reached
estimated maxima of 19.6, 2.3, and 12 per 1000 m? during 1977. Maximum densities of all species were found within 50 m of
the nearest water body in moist habitat vegetated by sedges, grasses, horsetails, willows, and poplar. Lower densities were
found in upland mixed woods and no anurans were found in dry areas with sandy substrate and Jack Pine forest. Other
amphibian and reptile species were not encountered.
Key Words: habitat, population densities, boreal forest, Alberta, Wood Frog (Rana sylvatica), Boreal Chorus Frog
( Pseudracis triseriata maculata), Canadian Toad (Bufo hemiophrys).
The herpetofauna of northeastern Alberta is
poorly known. Only a few locality records for
four species of amphibians and a single reptilian
species exist for northeastern Alberta (Logier
and Toner 1961). The Canadian Toad (Bufo
hemiophrys), the Boreal Chorus Frog ( Pseuda-
cris triseriata maculata), and the Wood Frog
(Rana sylvatica) are known from the Fort
McMurray area (University of Alberta Museum
of Zoology = UAMZ collections). Little has
been published regarding the abundance and life
histories of these species within Alberta, especi-
ally in the northern half of the province. Harper
(1931) reports the occurrence of the Leopard
Frog (Rana pipiens) and Red-sided Garter
Snake (Thamnophis sirtalis parietalis) in the
extreme northeastern corner of Alberta. The
present study was undertaken to determine
reproductive phenology, habitat utilization, and
population densities of the three amphibian
species commonly found in the boreal forest
region of northeastern Alberta.
Most amphibians and reptiles appear to have
well defined home ranges (Porter 1972). Quanti-
tative studies of home ranges have been con-
ducted for the Wood Frog by Bellis (1965), the
Chorus Frog by Kramer (1973, 1974), and the
Canadian Toad by Breckenridge and Tester
(1961). The tendency for individuals to spend
extended periods of time at a given locality
permits population estimates to be made by
“total” counts, that is, the sum of individuals
marked (counted) within a given area. Such
counts may be higher than the actual number
present within an area at a given time. This
overestimate is the result of (1) counting all (or
nearly all) true residents plus a number of
transients; (2) counting individuals whose home
ranges do not lie completely within the study
plots; (3) mortality not being taken into account.
The studies by Breckenridge and Tester
(1961), Kramer (1973, 1974), and Bellis (1965)
were conducted in relatively small areas and thus
provide a limited amount of comparative infor-
mation with respect to habitat preferences.
Information on habitat utilization can be ob-
tained by conducting regular periodic censuses
of a large number of sample sites selected to
include a wide range of habitat types.
Study Area
Twenty-four potential spawning sites (Figure
1) were selected in 1976 and 1977 within the
boreal forest region of northestern Alberta.
These sites consisted of 12 streams, five borrow
pits (excavations resulting from the removal of
144
1979
ROBERTS AND LEWIN: AMPHIBIAN POPULATIONS. ALBERTA
Peace River Lake
oo Athabasca
Claire
Wood Buffalo
National
Athabasca
River
Edmonton,
520 miles
20 30 km
Birch sete:
Mountains
Ells River
Gregoire
\
\ J SS
t
“4
6
8 9
loose
Ss.
oe
+
rks ¢/ HZ
FIGURE |. Location of amphibian study sites in northeastern Alberta.
145
146
road building materials), four natural ponds,
portions of the shores of two lakes, and a bog.
During 1977, 16 of these sites plus a dry Jack
Pine (Pinus banksiana) ridge were studied to
determine habitat utilization and population
density.
Methods
Each site was visited during the day at least
eight times during June, July, and August and
the amphibian species present, and their abun-
dance, were recorded in the following manner.
The visits were equally distributed during morn-
ings and afternoons and as well between cool
and warm days for each site. During each visit
the observer walked completely around the
margins of ponds and borrow pits or fora fixed
distance (approximately 400 m) along the mar-
gins of lakes or flowing waters. Amphibians .
observed were identified to species and the
numbers of individuals of each were recorded.
Individuals were assigned to developmental
stages (if larvae) or size classes (if terrestrial) in
order to describe their development, time of
metamorphosis, and growth during their first
year of life. The following arbitrary classes were
employed, with equivalent stages of Gosner
(1960) given in brackets: (1) eggs (1-25); (2)
tadpoles without limbs (26-31); (3) tadpoles with
hind limbs present (32-40); (4) tadpoles with
four limbs present (41-42); (5) transforming
tadpoles (semi-aquatic or terrestrial with tail still
evident) (43-45); (6) young-of-the-year with tail
completely resorbed (46); (7) yearlings (the
smallest size class of subadults or adults); (8)
adults (large subadults and adults). The smaller
number of classes or stages distinguished by
relatively gross differences facilitates the des-
cribing of large numbers of larvae in the field.
The head-body lengths (= length) of post-
metamorphic frogs and toads were measured to
the nearest 0.1 mm according to Conant (1975).
Small subadults were similarly measured, up to
and including the time of metamorphosis of
young-of-the-year. Young-of-the-year were
measured again in late August. The data should
provide objective criteria for distinguishing
underyearling (class 6), yearling (class 7), and
older (class 8) individuals, as preliminary obser-
vations here indicate that these groups occur in
mutually exclusive length-frequency groups.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Subsamples of aggregations of larvae were
collected and preserved in 10% formalin. These
were later identified to genus using keys in Blair
et al. (1968). Specimens of each species and their
developmental stages from each site were col-
lected throughout the study and deposited inthe
University of Alberta Museum of Zoology
(Accession number 76-31).
At each habitat sampling site three plots,
10 X 100 m, were established at right angles to
the water body (if present). Where more than
one habitat type was present the plots were
established so that all types were sampled.
Where a water body was present the following
information was recorded: basin morphology,
submergent and emergent vegetation present.
Owing to the large area encompassed by the
sample plots only the dominant vegetation types
were mapped for each plot.
A census of each of the sample plots was taken
10 times during the summer of 1977. During
each visit the observer noted weather conditions,
walked the plot, traversing its width 20 times
going away from the pond edge and 20 times on
the return trip to the pond ina crisscross pattern.
During each visit the species present, age class
(as determined by length), location on the plot,
and habitat type with which each individual was
associated was recorded. Subadults and adults
were marked by removal of the phalanges distal
to the web on one digit of the left hind foot: thus
they were not individually marked. The popula-
tion of adults on each plot was regarded as the
total number of individuals so marked.
The sample plots were subdivided by orange
surveyor’s tape placed at 20-m intervals. Indi-
viduals found between the water margin and the
20-m marker were assigned the mean value of
10 m, and individuals found between 20- and 40-
m markers were assigned the mean value of 30 m
and so on for purposes of calculating the mean
distance that each species was found from the
water margin during each month of the study.
Results and Discussion
During the 1976 survey 478 adult and sub-
adult amphibians were counted of which 225
were Wood Frogs, 207 Canadian Toads, and 16
Boreal Chorus Frogs. No spawning pairs or egg
masses were found during the present study.
IY)
Tadpoles were present in all of the known
spawning sites during mid-June.
Abundance, Spawning, and Development
Wood Frog
The Wood Frog was present at all sites and
spawned in 14 of the 24 sites studied. This species
spawned in natural ponds, borrow pits, and a
backwater on a stream. Egg masses of this
species are attached to aquatic vegetation
(Wright and Wright 1949): thus Wood Frogs are
most likely to spawn in standing water where
suitable vegetation may be found. Spawning had
apparently been completed before the 1976
study was initiated in mid-June. This species
spawns as early as Marchin the southern portion
of its range but as late as July in the Northwest
Territories (Wright and Wright 1949). Within
the study area metamorphosis occurred as early
as 30 June and was complete by early August.
The period during which metamorphosis occur-
red within the study area was exceptionally
short. Metamorphosis elsewhere may occur
from late May to mid-September (Wright and
Wright 1949) and overwintering of tadpoles may
occur at northern latitudes (Bleakney 1954).
Wood Frogs metamorphose at a size of
14-22 mm_ head-body length (Wright and
Wright 1949). Postmetamorphic individuals in
the present study ranged from 14.4 to 18.7 mm
CsI pe NE= 918) and trom: se) ton24-0, mim
(x = 20.5, N=12) by late August. Yearlings
ranged from 20.4 to 31.0 mm (x = 25.4, N = 13)
in late June and thus can be readily dis-
tinguished from young-of-the-year. Yearlings
apparently do not spawn, as frogs of this size
were not found in breeding aggregations. The
length of spawning individuals ranges from 29 to
50 mm for males and 34 to 56 for females
throughout the range of this species (Wright and
Wright 1949). Individuals of this size within the
study area are almost certainly two or more
years old. Schueler (1973) also found that frogs
of this species did not breed until they were two
or more years old.
Dispersal of post-spawning adults away from
the spawning sites occurs but some adults may
be encountered near the margins of water bodies
throughout the summer. Young-of-the-year also
leave the spawning sites shortly after meta-
morphosis and only small numbers were en-
ROBERTS AND LEWIN: AMPHIBIAN POPULATIONS, ALBERTA 147
countered at these sites during late August.
Wood Frogs are terrestrial and are occasionally
found far from water in moist wooded areas.
They are known to overwinter under leaf litter
(Wright and Wright 1949; Hodge 1976) and
under logs or stumps.
Boreal Chorus Frog
The Boreal Chorus Frog was seen at seven of
the study sites and spawned at three of these.
This species spawned in two of the borrow pits
studied and in one natural pond. The eggs are
laid in small loose clusters attached to vege-
tation, frequently in temporary ponds (Wright
and Wright 1949). Spawning within the study
area had been completed prior to 12 June 1976.
Spawning is known to occur as early as late
March in Indiana (Wright and Wright 1949),
and as early as late April at Lake Wabamun,
Alberta. Calling individuals have been heard as
early as 3 April (1976) along the Red Deer River
near Innisfail (52°N) (personal observations
W.R.). Spawning was observed in the north-
western quarter of Alberta in late May of 1976
and likely occurred within the study area at that
time. Metamorphosis occurred as early as 24
July and was completed shortly after 3 August.
As with the Wood Frog, metamorphosis of this
species occurred within a short period. Boreal
Chorus Frogs throughout their range meta-
morphose at lengths of 7.5-13.0 mm (Wright
and Wright 1949). Postmetamorphic individuals
measured during the present study were much
larger, 12.6-14.8 mm (x = 13.7, N = 14). Other
northern Alberta specimens (N = 18, UAMZ
collection) averaged 12.7 mmat, or shortly after, |
metamorphosis. The relatively large size of
Boreal Chorus Frogs at the time of metamor-
phosis may be the result of development at low
temperatures as is true for many poikilotherms
(Ray 1961). The size at which young-of-the-year
overwinter may be a critical factor affecting their
ability to persist during periods of stress. Larger
individuals may be more successful in this
regard. Yearlings in a number of northern
Alberta collections ranged from 15.9 to 17.3 mm
during May. Spawning adults, taken from
breeding aggregations, in Northern Alberta
ranged from 19.3 to 28.2 mm (Xx = 23.0, N = 28).
The size of spawning individuals throughout the
range of this species ranges from 21 to 32 mm for
148
males and 20.0 to 37.5 mm for females (Wright
and Wright 1949) indicating that individuals
within the study area probably do not spawn
until they are two or more years old. It is of
interest that Matthews (1971) reports that
marked individuals in a montane population do
not breed until their fourth summer. Boreal
Chorus Frogs are known to overwinter in
Alberta by burrowing under decaying stumps
and into anthills (personal observations W. R.).
Canadian Toad
The Canadian Toad was present at nine of the
study sites during 1976 and spawned at seven of
these. This species utilized the widest variety of
spawning habitats including natural ponds,
borrow pits, streams, and lake margins. The eggs
of Canadian Toads are laid in long strings and
may or may not be associated with vegetation
(personal observations W. R.). The long strings
of eggs may be less subject to displacement by
moving water than the attached spherical or
subspherical egg masses of frogs and thus
facilitate spawning in waters with a slight
current, wave action, and no vegetation. This
increases the variety of spawning habitats
available to the species; however, egg and
tadpole mortality may be higher owing to
predation and displacement by running water.
The fecundity of Canadian Toads (3354-5842
eggs, N =3 (V. L. unpublished data)) is higher
than that of Wood Frogs (x = 777 eggs (Herreid
and Kinney 1966)) and Chorus Frogs (x = 597
eggs (Whitaker 1971)), which may compensate
for the mortality arising in the wide variety of
spawning habitats utilized. These strings are also
adaptive as some eggs will usually remain
submerged if the water level recedes. Spawning
within the study area had apparently been
completed prior to 12 June. Spawning com-
menced during May along the Red Deer Riverin
1976 and had occurred into mid-June at the
Same site in previous years. Within the study
area metamorphosis occurred as early as 25 June
and continued until early July. The develop-
mental time for this species appears to be
unknown. Postmetamorphic individuals of this
Species range from 9.0 to 13.5 mm in head-body
length (Wright and Wright 1949). Individuals
sampled during the present study ranged from
IS*to 13.2 mm (x = 1274- N= 11) and had at-
tained head-body lengths of 19.3-28.0 mm
THE CANADIAN FIELD-NATURALIST
Vol. 93
(xX = 24.8, N = 10) by late August. The smallest
yearling found in late June was 22.0 mm long;
thus yearlings are readily distinguishable from
young-of-the-year. Spawning individuals of this
species range from 58 to 68 mm for males and 56
to 80 mm for females (Wright and Wright 1949):
it is unlikely that individuals of this species
within the study area spawn until they are two or
more years old.
Dispersal of adults from spawning sites occurs
soon after spawning but individuals may be
found along the margins of these sites through-
out the summer. Young-of-the-year toads are
often found in high numbers along the margins
of spawning areas throughout the summer.
Lillywhite and Wassersug (1974) regard the
aggregating behavior of young-of-the-year Bor-
eal Toads (Bufo boreas) as a retained larval
characteristic. Young-of-the-year Canadian
Toads were abundant along the Red Deer River
(a spawning site) in early September of 1976 but
were absent after mid-September, probably
having buried themselves in suitable substrate
for overwintering.
Habitats and Population Densities
Nine habitat types were identified within 17
study sites. These are ranked in order of
decreasing wetness as follows: (1) water, (2)
sedge (Carex spp.) fen, (3) grass (Graminae)
meadows, (4) willow (Salix spp.) bog, (5)
Trembling Aspen (Populus tremuloides) forest,
(6) Black Spruce (Picea mariana), (7) White
Spruce (Pica glauca), (8) upland mixed wood
(Populus tremuloides, Picea mariana, _ P.
glauca), (9) Jack Pine (Pinus banksiana).
Because visits to each site were equally divided
between mornings and afternoons and pro-
portionately between cool and warm days, any
biases in the number of amphibians observed
resulting from these factors were minimized. In
fact 28.6% of the amphibians were counted on
cool mornings (23.7% of the visits) and 14.4%
were seen on cool afternoons (14.8%): 34.7%
were seen on warm mornings (33.1%), and 22.3%
were seen on warm afternoons (28.4%).
Wood Frog
The extent to which each habitat type was
utilized by each age class of the Wood Frog
during May—August of 1977 is shown in Figure
2. Wood Frogs were most frequently found in
1979
KEY
HABITAT TYPES:
| Water
2 Sedge Fen
3 Grass Meadow
4 Willow-bog
5 Aspen Poplar
6 Black Spruce
7 White Spruce
8 Upland Mixed Wood
9 Jack Pine Dry
SYMBOLS}
15
| | Adults (N=15)
10
Ne Yearlings (N=10)
Z Young-of-year (N=5)
rn)
Ww
a
=
5)
2
z
=
2
=
a
2
<
5
O
ROBERTS AND LEWIN: AMPHIBIAN POPULATIONS, ALBERTA 149
WOOD FROG
FIGURE 2. Habitat utilization by Wood Frogs in northeastern Alberta during May—August 1977.
grass meadows, willow bogs, and aspen poplar
habitat, which are the most moist terrestrial
habitats available. Heatewole (1961) regards the
Wood Frog as a forest-dwelling species and
suggests that the shade provided by trees is an
important factor in determining habitat selec-
tion. He demonstrated experimentally that
substrate structure and substrate moisture are
important factors governing the selection of
microhabitat. Bellis (1962) states that Wood
Frogs are restricted to forested areas, particu-
larly lowland bogs. Shelford (1913) demon-
strated that Wood Frogs prefer an environment
with moist air. Marshall and Buell (1955) found
that in exposed areas with sparse Wood Frog
populations the vapor pressure deficit was
higher than in shaded tamarack (Larix /arcinia)
habitat with a dense Wood Frog population. It
thus appears that moisture content of both the
substrate and air are important factors in
determining the suitability of habitat types for
Wood Frogs. Tall grasses, willows, and aspen
poplars growing in moist soil in low-lying areas
provide a favorable environment for Wood
Frogs within the study area, while much of the
forest habitat, especially Jack Pine area, is too
dry with respect to substrate moisture and
perhaps the humidity of the air.
Heatwole (1961) suggests that the Wood Frog
is less terrestrial than previously thought (see
Wright and Wright 1949) and points out that its
association with woods and damp leaf litter
occurs after woodland ponds dry up. Our study
sites were associated with more permanent
bodies of water. Wood Frogs were rarely found
at distances greater than 100 m from a water
body and the mean distance for all age classes
during each month was less than 50 m. Bellis
150
(1962) regards the Wood Frog as intermediate
between aquatic and terrestrial anurans in its
ability to withstand body water loss. Only large
Wood Frogs are found in dry habitats and it is
probably the large individuals that emigrate and
colonize new areas (Bellis 1962). We also noted
that larger Wood Frogs tended to be further
from water, perhaps owing to their greater
ability to tolerate body water loss.
During 1977, 646 Wood Frogs were counted.
Population densities of this species ranged from
0.0 to 19.6 individuals (yearlings or older) per
1000 m2 (Table 1). The highest population
density for this species (19.6 adults per 1000 m2?)
was found in an area with a mixture of grasses,
willow, and aspen poplar. These plant communi-
ties are frequently found individually, mixed or
intergrading with one another adjacent to water
bodies. Bellis (1961) found maximum densities
of about 17 individuals per 1000 m2 in Tamarack
and Calla Lily (Calla palustris) habitat.
Boreal Chorus Frog
Boreal Chorus Frogs were most frequently
found near the water margin or in shallow water
amongst vegetation (Figure 3). Too few year-
lings or adults were found to determine what, if
any, habitat type is preferred. Whitaker (1971)
regards the Chorus Frog as a woodland species
that occurs most frequently near water, while
Stebbins (1954) notes that in the west this species
seems to be primarily an inhabitant of open
moist grasslands. Kramer (1974) found that the
TABLE |—Densities of amphibians in selected habitat types
within the study area. Numbers in parentheses indicate
maximum estimates
Density per 1000 m2
Boreal
Habitat Wood Chorus Canadian
types Frog Frog Toad
Water
Sedge
Willow (2.3) 0.45
Grass (19.6) 2.83 (12) 0.83
Horsetail
Poplar
White Spruce
Black Spruce 1.08 0.07 0.16
Upland mixed j
wood
Jack Pine 0 0 0
THE CANADIAN FIELD-NATURALIST
Vol. 93
home ranges of Chorus Frogs contained breed-
ing pools and that most frogs remained within
100 m of these pools throughout the summer. In
Alberta the Boreal Chorus Frog is known from
the prairies, aspen parkland, and boreal forest
regions (UAMZ collections), but always near
water. It is the most widespread amphibian
species within Alberta and persists in a wide
variety of moist habitat types providing that at
least temporary ponds are available for spawn-
ing and for development of the tadpoles.
Boreal Chorus Frogs were found closer to
water on the average than the other anuran
species present. Adults and young-of-the-year
were found at a mean distance of less than 30 m
from water during all months. During June of
1977 nonbreeding individuals were found at a
mean distance of 54 m from the water margin.
This may be the result of their being found close
to their overwintering sites. Unlike breeding
individuals they probably do not move immedi-
ately to water bodies after emergence in the
spring. During July and August most indi-
viduals of all age classes were found within 20 m
of the water margins. Kramer (1973) notes that
while a few individuals may be found over 200 m
from breeding pools, most are found within
100 m. Individuals may be found buried under
leaf litter during the day (Kramer 1973). This
may serve as protection against diurnal preda-
tors and desiccation. These observations may
indicate that this small species is more prone to
desiccation than are the larger species.
During 1977, 148 Boreal Chorus Frogs were
counted. Insufficient data were obtained to
determine population densities on most study
plots. Two sites yielded adult population densi-
ties of 2.3 individuals per 1000 m2 (Table 1).
Kramer (1974) obtained densities of 0.55 adults
per 1000 m2 from a 33-acre (13.4-ha) study area
in Indiana. His data probably include most of
the adults present; estimates from the present
study, on the other hand, are probably con-
servative owing to the secretive nature of the
Boreal Chorus Frog. Larger numbers of the
Boreal Chorus Frog are assumed to be present
because of the numbers heard calling (but
seldom seen) during May and June.
Canadian Toad
The Canadian Toad was found most fre-
quently in grass meadows and willow bogs near
1979
BOREAL CHORUS
KEY
HABITAT TYPES:
| Water Wet
2 Sedge Fen
3 Grass Meadow
4 Willow- bog
5 Aspen Poplar
6 Black Spruce
7 White Spruce
8 Upland Mixed Wood
9 Jack Pine Dry
SYMBOLS:
15
- | | Adults (N=15)
uy b:] Yearlings (N=10)
6 Z Young -of—year (N=5)
”
a
Ww
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Zz
z
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es)
ce
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123456789
MAY
ROBERTS AND LEWIN: AMPHIBIAN POPULATIONS, ALBERTA 15]
FROG
HABITAT TYPES
FiGuRE 3. Habitat utilization by Boreal Chorus Frogs in northeastern Alberta during May—August 1977.
the water’s edge (Figure 4). The Canadian Toad
was common at only three of the 17 sites studied.
Each of the three sites included poplar forest and
grass and two of the three included willows. It is
of note that each site was associated with a river
or lake rather than a temporary or small
permanent pond. Stebbins (1954) states that this
species occurs in Transition and lower Boreal life
zones (Transition and Canadian life zones of
Soper(1964)), in the vicinity of lakes and streams.
Underhill (1961) and Henrich (1968) regard the
Canadian Toad as a water-adapted species (1.e.,
less terrestrial than most bufonids) of prairie
ponds and lakes. Breckenridge and Tester (1961)
found that this was the only toad species present
in habitat vegetated by aspen, willows, bluestem
(Andropogon spp.), and grasses adjacent to a
lake. This species is not a forest inhabitant but
resides within damp open areas adjacent to
water bodies. Its association with lakes and
flowing water may be in response to its ability to
utilize them as spawning and rearing habitats. In
such habitats this species is subject to a
minimum of competitive interactions with other .
anurans which are unable to reproduce success-
fully there.
Canadian Toads are seldom found far from
water. Breckenridge and Tester (1961) found
that only a few toads were captured in a trap
63.3 m (200 ft) from the water margin while over
80% of the total capture was within 8 m (25 ft).
In June the mean distance from water for all age
classes was 60 m; in July and August the mean
distance for all age classes was less than 50 m.
This difference may have been in response to
drying of the terrestrial habitat as the summer
progressed. Tamsitt (1962) reports migration of
this species to water margins at night, pre-
sumably for water imbibition. Breckenridge and |
Tester (1961) found no such nightly migration,
152 THE CANADIAN FIELD-NATURALIST
Keay
HABITAT TYPES:
| Water
2 Sedge Fen
3Grass Meadow
4 Willow-bog
5 Aspen Poplar
6 Black Spruce
7 White Spruce
8 Upland Mixed Wood
9 Jack Pine Dry
SYMBOLS}
n
a
WwW
a
=
>)
z
z
=<
=
x
a
=
<=
2308 6 7B ©
M AY
Vol. 93
CANADIAN TOAD
Bernd DOQQ
iS 73 ©
AUGUST
HABITAT TYPES
FIGURE 4. Habitat utilization by Canadian Toads in northeastern Alberta during May—August 1977.
however; individuals were seldom found suf-
ficiently far upland that they were in particularly
dry habitat.
During 1977, 130 Canadian Toads were
counted. Population estimates for this species
reached a maximum of 12 adults per 1000 m2
(Table 1), only slightly over half of the maximum
density for Wood Frogs. Kelleher and Tester
(1969) found Canadian Toads in much greater
abundance. This was probably owing to an
abundance of excellent overwintering habitat
combined with a lack of competition from other
anurans along the margins of a prairie lake.
General Discussion
The relative abundance of the three species
found during both years of the study may be
misleading. The Wood Frog is certainly wide-
spread and abundant within the study area. This
species is the most widely distributed amphibian
in North America and occurs further north than
any other species. Its adaptations to life in cold
regions are summarized by Hodge (1976).
Canadian Toads, however, may be much less
abundant than the Wood Frog, as 182 of the 207
adult and subadult Canadian Toads seen during
1976 were young-of-the-year. These were found
in postmetamorphic aggregations along the mar-
gins of spawning sites. In spite of its ability to use
the widest variety of spawning habitats (among
the three species found here) this species spawn-
ed in only 7 of the 24 sites studied compared with
14 sites utilized by the Wood Frog. The Boreal
Chorus Frog was heard at or near all of the study
sites but spawned in only three of these. Its
utilization of temporary ponds and standing
bodies of water for spawning sites may reduce
the number of potential sites from 24 to 10 for
this species. The small size, cryptic coloration,
and secretive behavior of Boreal Chorus Frogs
1979
ROBERTS AND LEWIN: AMPHIBIAN POPULATIONS, ALBERTA 153
CANADIAN TOAD
2
1000m
9
ine)
::Subadults: 2222)
°
Individuals /
(eo)
Wiilow
Sedge, Grass
=
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ac
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Oo
a.
Horsetail
Cattail
60
CHORUS FROG
WOOD FROG
Upland Jack Pine
Mixed
Sand Ridge
DISTANCE FROM WATERIN METRES
FIGURE 5. Schematic distribution and population density of adult and subadult amphibians in northeastern Alberta in
relation to vegetation and water.
make them difficult to see and collect even when
they are present in large numbers at spawning
SiLeSee SINCE teas) sae stenhesthial species. and
frequently hides under logs and stones, it is the
most difficult to see of the three species present.
This species 1s probably more abundant than our
data indicate.
Leopard Frogs are apparently absent from
our study area although they occur further
north, probably as glacial relicts. The Red-sided
Garter Snake was also not encountered; how-
ever, it may occur sporadically, as we have a
recent and reliable sight record of one at Point de
Roche on the west shore of Lake Claire (J.
Kristensen, Department of Zoology, University
of Alberta, personal communication).
Thus the known herpetofauna of northeastern
Alberta consists of four amphibian species: the
Leopard Frog, Wood Frog, Boreal Chorus
Frog, and Canadian Toad, and a single reptile,
the Red-sided Garter Snake. The Wood Frog is
the most abundant and widespread species. The
three common amphibian species share a num-
ber of life history features. They overwinter in
terrestrial sites and spawn in the spring. Larval
development is rapid and metamorphosis occurs
during the first summer of life. Borrow pits as
well as natural ponds appear to be favored as
spawning sites over streams and lakes, although
lakes are used by toads. Subadults and adults are
terrestrial; however, most are found within 50 m
of a water body. Highest densities of all species
are found in moist habitats, near water vegetated
by sedges, grasses, horsetail, willow, and aspen.
The habitat utilization, population density,
and distribution of age classes for each species is
schematically summarized in Figure 5. The
vegetation profile is based on the frequency of
occurrence of dominant species within each
20-m segment of the 48 sample plots adjacent to
154 THE CANADIAN FIELD-NATURALIST
water. The density values for each amphibian
species are based on totals of subadults and
adults for all sites and thus represent mean
densities. Wood Frogs may be found over 100 m
from water; however, most are found within
50 m. Small individuals tend to be found closer
to water than large ones. Chorus Frogs are
uncommon at distances greater than 20 m from
water and none were found further than 100 m
- from water. Canadian Toads gradually decrease
in abundance at distances greater than 40 m
from water. No amphibians were found in dry
sandy areas with Jack Pine forest habitat.
Acknowledgments
Financial support for this study was provided
by Alberta Oil Sands Environmental Research
Program (AOSERP Project TF 5.1). L. Brusnyk
provided excellent field assistance and, as well,
helped with data analysis. Both L. Brusnyk and
B. Herbert prepared the figures. J. O. Murie
provided useful suggestions during the initial
phase of this study.
Literature Cited
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wood frog in a Minnesota bog. Herpetologica 17: 228-
231.
Bellis, E.D. 1962. The influence of humidity on wood
frog activity. American Midland Naturalist 68: 139-148.
Bellis, E. D. 1965. Home range and movements of the
wood frog in a northern bog. Ecology 46: 90-98.
Blair, W.F., A. P. Blair, P. Brodkorb, F. R. Cagle, and
G. A. Moore. 1968. Vertebrates of the United States.
2nd edition. McGraw-Hill Book Co., New York. 616 pp.
Bleakney, J. S. 1954. Range extensions of amphibians in
eastern Canada. Canadian Field-Naturalist 66: 165-171.
Breckenridge, W. J. and J. R. Tester. 1961. Growth, local
movements and hibernation of the Manitoba toad, Bufo
hemiophrys. Ecology 42: 637-646.
Conant, R. 1975. A field guide to reptiles and amphibians
of eastern and central North America. 2nd edition.
Houghton Mifflin Co., Boston. 429 pp.
Gosner, K. L. 1960. A simplified table for staging anuran
embryos and larvae with notes on identification. Herpe-
tologica 16: 183-190.
Harper, F. 1931. Amphibians and reptiles of the Athabasca
and Great Slave lakes region. Canadian Field-Naturalist
45: 68-70.
Heatewole, H. 1961. Habitat selection and activity of the
wood frog, Rana sylvatica Le Conte. American Midland
Naturalist 66: 301-313.
Henrich, T. W. 1968. Morphological evidence of secon-
dary intergradation between Bufo hemiophrys Cope and
Bufo americanus Holbrook in eastern South Dakota.
Herpetologica 24: 1-13.
Vol. 93
Herreid, C. F. and S. Kinney. 1966. Survival of Alaskan
Woodfrog (Rana sylvatica) larvae. Ecology 47: 1039-
1041.
Hodge, R. P. 1976. Amphibians and reptiles in Alaska, the
Yukon and Northwest Territories. Alaska Northwest
Publishing Co., Anchorage, Alaska. 89 pp.
Kelleher, K. E. and J. R. Tester. 1969. Homing and sur-
vival in the Manitoba toad, Bufo hemiophrys, in Min-
nesota. Ecology 50: 1040-1048.
Kramer, D. C. 1973. Movements of western chorus frogs
Pseudacris triseriata triseriata tagged with Co. Journal
of Herpetology 7: 231-235.
Kramer, D. C. 1974. Home range of the western chorus
frog Pseudacris triseriata triseriata. Journal of Herpe-
tology 8: 245-246.
Lillywhite, H. B. and J. Wassersug. 1974. Comments on a
postmetamorphic aggregate of Bufo boreas. Copeia 1974:
984-986.
Logier, E.B.S. and G.C. Toner. 1961. Check list of
amphibians and reptiles of Canada and Alaska, a
revision of contribution No. 41. Royal Ontario Museum,
Contribution Number 53. 92 pp.
Marshall, W.H. and M. F. Buell. 1955. A study of the
occurrence of amphibians in relation to a bog suc-
cession, Itasca State Park, Minnesota. Ecology 36: 381—
387.
Matthews, T. C. 1971. Genetic changes in a population
of boreal chorus frogs ( Pseudacris triseriata) polymorphic
for color. American Midland Naturalist 85: 208-221.
Porter, K.R. 1972. Herpetology. W.B. Saunders Co.,
Toronto. 523 pp.
Ray, C. 1961. The application of Bergmann’s and Allen’s
rules to the poikilotherms. Journal of Morphology 106:
85-108.
Schueler, F. W. 1973. Frogs of the Ontario coast of
Hudson Bay and James Bay. Canadian Field-Naturalist
87: 409-418.
Shelford, V. E. 1913. The reactions of certain animals to
gradients of evaporating power of air. A study of
experimental ecology. Biological Bulletin (Woods Hole)
25: 79-120.
Soper, J.D. 1964. The mammals of Alberta. Hamly Press
Ltd., Edmonton. 402 pp.
Stebbins, R. C. 1954. Amphibians and reptiles of western
North America. McGraw-Hill Book Co., Toronto. 536 pp.
Tamsitt, J. R. 1962. Notes ona population of the Manitoba
toad (Bufo hemiophrys) in the Delta marsh region of
Lake Manitoba, Canada. Ecology 43: 147-150.
Underhill, J. C. 1961. Intraspecific variation in the Dakota
toad, Bufo hemiophrys, from northeastern South Dakota.
Herpetologica 17: 220-227.
Whitaker, J.O. 1971. A study of the western chorus frog,
Pseudacris triseriata, in Virgo County, Indiana. Journal
of Herpetology 5: 127-150.
Wright, A. H. and A. A. Wright. 1949. Handbook of frogs
and toads of the United States and Canada. Comstock
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Received 8 August 1978
Accepted 27 November 1978
Caribou Distribution and Group Composition
Associated with Construction of the Trans-Alaska
Pipeline
RAYMOND D. CAMERON,! KENNETH R. WHITTEN, !
WALTER T. SMITH,2 and DANIEL D. ROBY:
‘Alaska Department of Fish and Game, 1300 College Road, Fairbanks, Alaska 99701
2Department of Biology, Idaho State University, Pocatello, Idaho 83209
3Alaska Cooperative Wildlife Research Unit, University of Alaska, Fairbanks, Alaska 99701
Cameron, Raymond D., Kenneth R. Whitten, Walter T. Smith, and Daniel D. Roby. 1979. Caribou distribution and group
composition associated with construction of the Trans-Alaska Pipeline. Canadian Field-Naturalist 93(2): 155-162.
Caribou surveys were conducted periodically along the Trans-Alaska Pipeline haul road on the central Arctic Slope between
June and November 1975. Mean calf percentage observed in summer was approximately one-third lower than that obtained
from concurrent aerial surveys of both the pipeline corridor and adjacent areas; however, fall means were identical. In both
summer and early fall, mean latitudes calculated for groups with and without calves along the haul road were more southerly
than for the corresponding group types observed through aerial survey. A more detailed regional comparison of survey data
revealed corridor-related abnormalities in Caribou distribution and group composition. No Caribou, or relatively low
numbers, were observed in the northernmost segment of the pipeline corridor near Prudhoe Bay, and calf percentages in
summer were consistently lower for each of four arbitrarily established regions of the haul road than expected on the basis of
aerial survey results; fall calf percentages did not differ appreciably. Mean group size was generally lower along the haul road
than for comparable areas to the east and west. Responses of Caribou to the pipeline corridor in general, and of cows and
calves in particular, are discussed in relation to inherent avoidance tendencies as modified seasonally by terrain, group
dominance, and human activity.
Key Words: Caribou, Rangifer, pipeline, disturbance, group composition, Prudhoe Bay, ecological distribution.
The discovery of vast petroleum reserves at tion-related human activity. Consequently, a
Prudhoe Bay in 1968 resulted in construction of | major study was undertaken to evaluate the
the Trans-Alaska Pipeline, designed totransport reactions of Caribou to the Trans-Alaska Pipe-
crude oil to Valdez on the Gulf of Alaska. North line on a seasonal basis and to document any
of the Yukon River, the pipeline and its associated shifts in annual movement patterns
associated haul road were to traverse the ranges and range occupancy. To date, this investigation
of two major subpopulations of Barren-ground _ has been largely restricted to the Arctic Slope.
Caribou (Rangifer tarandus granti), the Porcu- Based on summer and fall surveys in this
pine and Western Arctic herds (Skoog 1968; region between 1969 and 1972, A. Gavin (1973,
Hemming 1971), and concern was focused onthe Report by Atlantic Richfield Company) re-
potential restriction of Caribou movements. ported a decline in Caribou numbers from 26000
In 1971 and 1972 K. N. Child (1973, Comple- to 2500, and he thought these Caribou to be
tion Report, Alaska Cooperative Wildlife Re- “offshoots” from the Porcupine and Western
search Unit, University of Alaska), using Arctic herds. Cameron and Whitten (1979),
simulated pipeline and pipeline crossing struc- however, have recently identified a third sub-
tures at Prudhoe Bay, attempted to assess the population of about 5000 Caribou whose range
effectiveness of various above-ground designsin is centered on the pipeline route along the
permitting free movements of Caribou. The Sagavanirktok River (Figure |). Seasonal move-
majority of approaching Caribou avoided the ments are primarily north-south, between
simulation and, of the successful crossings, calving grounds near the coast and winter range
about 3 times as many were by use oframpsthan in the northern foothills of the Brooks Range.
by passage under the structure. This detailed The identity of this “Central Arctic herd” has
study provided valuable preliminary data but apparently been confounded by periodic influx
was limited to observations on summer range. from one or both of the larger adjacent herds,
Thus, little was known of the seasonality of although such outside influence was thought to
Caribou responses to pipelines and construc- be minimal during the course of the present
155
THE CANADIAN FIELD-NATURALIST Vol. 93
156
OLIKTOK |
ry 4
STATION
1
DEADHORSEC}/
» /
i”
Y
=
MITIIN LI
le!
le\ FRANKLIN
s) BLUFFS
PUMP STATION 2
KEY
PUMP. STATION 3 ee soe
—— Above-ground Pipeline
ecoeeeBuried Pipeline
= Survey Coverage
GALBRAITH-+
LAKE t
PUMP STATION 4
FIGURE |. Survey coverage along the Trans-Alaska Pipeline corridor and regional boundaries established for comparison of
haul road and aerial survey results.
IST)
study. This report deals with changes in lati-
tudinal position, group size, and composition of
Caribou along the Trans-Alaska Pipeline haul
road during summer and fall 1975. These data
are compared with similar values obtained
through concurrent aerial survey of alargerarea
extending east and west of the corridor.
The haul road between the Yukon River and
Prudhoe Bay was completed in fall 1974, and
pipeline construction efforts were near maxi-
mum during the major portion of this investiga-
tion. The pipeline is 122 cm in diameter and is
alternately buried and above ground depending
on soil stability; ground clearances of elevated
sections range from less than | m toa maximum
of about 5 m.
Methods
Systematic surveys were conducted from the
Trans-Alaska Pipeline haul road between June
and November 1975. Generally these were
scheduled twice-monthly, each survey consisting
of two complete trips over the 263-km distance
between Pump Station 4 and Prudhoe Bay
(Figure 1). As only half of the route could be
covered in | d, most surveys required 4 d for
completion. Owing to inclement weather, survey
components were not necessarily on consecutive
CAMERON ET AL.: CARIBOU DISTRIBUTION NEAR PIPELINE
157
days, but an attempt was made to minimize the
time interval for each series. Inclusive dates are
given in Table 1.
A pickup truck, generally with one driver/
observer, was used for all road surveys. Speeds
were adjusted to 40-65 km/h, depending on
visibility, to provide reasonably constant per-
ceptibility of Caribou for anestimated 1.5 kmon
either side of the haul road. Only Caribou
observed initially with the naked eye were
recorded, but binoculars and spotting scope
were used as required to obtain additional
information. Pertinent data recorded for each
Caribou sighting included the following:
Location — road distance from a known point:
Number of Caribou per group ( defined arbi-
trarily as a single Caribou, or two or more
Caribou separated by less than an estimated
300 m);
Group composition —
calf — less than | yr old,
yearling — more than | yr old but less than
2 yr old,
cow — female more than 2yr of age,
bull — male more than 2 yr of age,
adult — more than | yr old, sex unknown,
unknown — unclassified as to sex or age.
TABLE I—Changes in Caribou numbers and mean group composition along the Trans-Alaska Pipeline haul road,
June—November 1975
Groups without
Total classified Groups with calves calves
Total number Number Number Number
Survey of caribou of Bulls, Calves, of Bulls, Calves, of Bulls,
dates observed caribou % % caribou” % % caribou” %
June 11-18 91 79 66 0 0 0 0 79 66
June 24-July 2 361 342 86 3 19 8 21 323 92
July 10-17 677 677 52 32 499 35 43 178 100
July 24-August 2 136 136 98 l 5 60 20 131 100
August 7-13 273 273 TH 7 Sy) 2 32 216 97
August 20-28 156 149 92 l 11 19 18 138 98
September 3-6 202 200 83 3 23 30 26 177 90
Summer mean*® 65 13(21) 31 40(35) 94
September 20-28 602 416 44 17 333 30 23 83 99
October 24-28 54 31 36 24 25 26 30 6 80
November 5-10 176 37 49 10 19 25 19 18 74
November 19-25 92 36 43 11 25 30 17 11 73
Fall mean® 43 17(17) 29 23(21) 92
“Total caribou in groups with no “unknowns” (see Methods).
*Total caribou in groups with one or more calves present.
“Total caribou in groups with no calves.
“Aggregate percentages; those in parentheses were determined from aerial surveys (Cameron and Whitten 1979).
158
Road locations were converted to their lati-
tudinal equivalents using 1:63 360 scale maps of
USGS Topography Series. For each survey
series a mean latitudinal position of Caribou was
calculated as described by Cameron and
Whitten (1979).
Results and Discussion
Caribou Numbers and Group Composition
Numbers and group composition of Caribou
observed from the haul road during each survey
period are given in Table |. Groups with one or
more “unknowns” (see Methods) were excluded
from composition calculations. Ninety-eight
percent of the total Caribou sighted during
summer were successfully classified. During fall,
however, bright sun and snow frequently created
distortion which precluded classification at
distances greater than about 400 m, and entire
groups were recorded as unknowns; the pro-
portion classified decreased to 56%. Cows and
yearlings were often difficult to distinguish, and
many such individuals were classified as “adults”
(see Methods) when more specific identification
was impossible. Calves, however, were relatively
easy to recognize and reported percentages are
thought to be very reliable. Similarly, iden-
tification of bulls was rarely in question.
During summer 13% of total Caribou ob-
served from the haul road were calves, com-
pared with a corresponding value of 21%
obtained by systematic aerial survey of a larger
area roughly centered on the pipeline route
(Table 1). In contrast, a mean of 17% calves was
observed in fall during both road and aerial
surveys, indicating that a representative portion
of the herd was present along the corridor.
Within each season, calf percentages for groups
with calves were similar for road and aerial
observations (Table 1). It can be calculated from
the summer road data in Table |, however, that
the number of Caribou observed in groups with
calves averaged 33% of the total classified,
whereas 60% of Caribou classified by air were
observed in groups with calves (see Cameron
and Whitten 1979): respective mean values for
fall were similar at 77 and 81%. In addition,
observations from the haul road indicate that
bulls were present in low numbers in groups with
calves but consistently predominated in groups
without calves, and the percentages did not
THE CANADIAN FIELD-NATURALIST
Vol. 93
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—-—8 Caribou in Groups w/Calves
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Cc
ro)
®
=
Jun Jul
Aug Sep Oct Nov
Month
FIGURE 2. Seasonal changes in mean latitudinal position and
mean group size of Caribou along the haul road and a
comparison with corresponding results from aerial surveys.
differ appreciably with season for either group
type (Table 1). Clearly then, the low proportion
of calves recorded during haul road surveys in
summer was due primarily toa local reduction in
the relative number and/or size (see below) of
groups with calves rather than a lower per-
centage of calves in such groups.
Latitudinal Movements
Changes in mean latitudinal position of
Caribou along the haul road are depicted in
Figure 2a. Throughout summer, groups with
calves remained farther north than groups
without calves. A similar trend was obtained
from concurrent aerial surveys (Cameron and
Whitten 1979), and these results are plotted for
comparison. Thus, the more northerly location
of the mean shown for aggregate sightings by air
ISA)
is partially attributable to a greater proportion
of groups with calves (see previous section)
because they tend to occupy higher latitudes.
Positions determined by aerial survey for groups
both with and without calves, however, were
also farther north than those calculated from
road survey observations. Corresponding means
did not differ appreciably in September, but the
same general trend was again apparent in
November (Figure 2a), despite the fact that no
overall differences in calf percentage were
observed (Table 1). Thus, dissimilar composi-
tion does not fully account for the more
southerly distribution of Caribou along the
pipeline corridor. Rather, local abnormalities
are indicated.
Group Size
Two distinct peaks in group size are shown in
Figure 2b for haul road observations, one in
mid-July during post-calving aggregation and
another in September just prior to the rut.
Except for late July and early September, the
mean size of groups with calves was greater than
that for groups without calves. A further
comparison of these means with those deter-
mined from aerial surveys indicates that in June,
August, and September the size of correspond-
ing group types was higher for aerial than for
haul road surveys. A valid comparison is not
possible for November because of the extremely
small sample size (Table |). Because the same
| June
O
Rr oeias > |
August
Percent of
Total Caribou Observed
CAMERON ET AL.: CARIBOU DISTRIBUTION NEAR PIPELINE
Bik
159
criteria for group identification were used in
both survey procedures (see Methods), these
data suggest avoidance of the area by larger
groups, group fragmentation, and/or a de-
creased tendency for group coalescence near the
pipeline corridor.
Differences in Regional Distribution and Group
Composition
To permit a more accurate assessment of
latitudinal distribution and group composition
of Caribou, four regions were delineated along
the pipeline corridor for direct comparison of
aerial and road survey observations. These
regions were centered on the corridor, and each
was bounded arbitrarily by '4 degree of latitude
and 2 degrees of longitude (Figure 1). The
regional distribution of total Caribou obser-
vations for both aerial and road surveys is shown
in Figure 3, and applicable calf percentages are
presented in Table 2. It was assumed that aerial
survey results reflected the true distribution of
Caribou among the four regions and provided
representative data on composition within each
region. Corresponding data from the haul road,
obtained concurrently, are compared with these
“expected” values. As sample sizes were fre-
quently low, statistical analysis of differences in
group composition was not feasible, but some
overall and regional trends are apparent.
Figure 3 demonstrates a distinct north-south
gradient in regional density of Caribou for both
November
[] Air
September
“abl sf hi.
a ZvOrenG [PREZ aac,
Region
FiGurE 3. Regional distribution of Caribou (see Figure 1) determined from corresponding aerial and haul road surveys.
160 THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 2—Regional comparison of calf percentages determined from corresponding aerial and haul road surveys,
June-November 1975
Region | Region 2 Region 3 Region 4
Number Number Number Number
Survey of Calves, of Calves of Calves, of Calves,
Month* method* caribou” % caribou” % caribou” % caribou” %
June Air 4 0 65 2 159 10 231 27
Road 0 121 0 195 4 26 12
August Air 0 — 49 0 122 9 200 34
Road 4 0 37 0 183 3 49 27
September Air 32 0 174 21 18 DD 99 20
Road 59 6 125 9 wy) 25 0 =
November Air 238 7 256 17 101 27 57 44
Road 36 11 0 — 0 — 0 =
“Aerial surveys (Cameron and Whitten 1979) or haul road surveys within the regions shown in Figure 1.
"Number of caribou classified, i.e., total caribou in groups with no “unknowns” (see Methods).
June and August, with aerial sightings in region
4 representing 50 and 55% of the total, respec-
tively. In contrast, the majority of haul road
observations during summer were in region 3,
and relatively few Caribou were observed near
the coast. Calf percentages for regions 3 and 4 of
the haul road were appreciably lower than
expected, while few calves were noted in regions
1 and 2 from either road or air (Table 2). For all
regions combined, calf percentages determined
from aerial and haul road surveys, respectively,
were 17 and 3 in June, and 20 and 7 in August.
Thus, the more southerly distribution of Cari-
bou within the pipeline corridor in summer was
due in part to a general avoidance of northern
sections near Prudhoe Bay. In addition, aggre-
gate and regional calf percentages were con-
sistently lower than the expected values, indi-
cating a disproportionate avoidance of the
pipeline corridor by groups with calves. The
resulting preponderance of bulls (i.e., groups
without calves), by virtue of their latitudinal
preference (Figure 2a), contributed further to a
southward displacement of Caribou along the
haul road.
By late September a southward movement of
Caribou had occurred. Approximately half of
the total sightings from both air and haul road
were in region 2, but no Caribou were observed
along the corridor in region 4 where nearly one-
third of the total was observed during aerial
surveys (Figure 3). During both aerial and road
surveys the majority of Caribou were found in
groups with calves (Table 1), and mean group
size increased (Figure 2b) as pre-rut aggrega-
tions formed. Although aerial observations
indicated that groups without calves remained
substantially farther south (Figure 2a), such
differences are of little quantitative importance
as these Caribou represented less than 10% of
total sightings. Regional differences in calf
percentage demonstrate no consistent trend
(Table 2), but combining data for regions 1, 2,
and 3 results in an estimated 17% calves for both
air and road surveys. Thus, in September a
strong avoidance of the Prudhoe Bay area is
indicated, but sightings of Caribou elsewhere
along the haul road appear to reflect overall herd
composition, and disproportionate avoidance
by groups with calves was not apparent. In
November aerial surveys established that fewer
Caribou were occupying coastal areas. Most
Caribou within the corridor were in region 1,
and none were observed in regions 3 and 4
(Figure 3). Total haul road sightings in Novem-
ber were among the lowest recorded (Table 1),
and any conclusions regarding differences in calf
composition would be equivocal because of
small sample size. Nevertheless, combined data
for fall again indicate that a more representative
portion of the herd approached and was ob-
served from the haul road, although continued
avoidance of northern areas is indicated.
Relatively low numbers of Caribou near the
haul road in region 4 during summer and fall
may partially explain the small group size
observed within the pipeline corridor (Figure
2b). Groups with calves observed along the coast
9
by air were generally larger than comparable
inland groups, and their avoidance of the haul
road area would tend to depress the calculated
mean irrespective of other influences on group
size. Groups without calves, however, exhibited
a similar decrease in summer and a corridor-
related reduction in group size remains suspect.
Avoidance Behavior of Caribou
Avoidance of the Prudhoe Bay area was noted
throughout the course of this study. The coastal
region near Prudhoe Bay, recently the site of
increased construction and exploration activity,
was previously a portion of the calving grounds
for the Central Arctic herd. As late as 1972 K. N.
Child (1973, op. cit.) and A. Gavin (1973, op.
cit.) reported calving within or immediately
adjacent to the Prudhoe Bay complex. With
facilities expansion and continued human activi-
ty over the past 3 or 4 yr, local occupancy by
Caribou has generally decreased, and in 1975 we
observed no neo-natal calves from this northern-
most section of the haul road. The area was also
previously invaded by Caribou during annual
post-calving movements along the coast (Cam-
eron and Whitten 1979) and during oscillatory
movements to and from the coast in response to
changing insect density (Child 1973, op. cit.;
White et al. 1975). Evidence for these occur-
rences 1s still visible as Caribou trail systems and,
although movements within the Prudhoe com-
plex are still detectable, they are now mere
remnants of past activities. Post-calving and
insect-induced movements, however, still occur
elsewhere along the coast between the Colville
and Canning rivers, indicating avoidance of a
specific portion of summer range with continued
occupancy of adjacent regions. Disturbance-
related abandonment of range is thought to be a
gradual process, occurring with increasing
avoidance of adverse stimuli (Klein 1971; Calef
1974), and the recent history of changing
Caribou occupancy near Prudhoe Bay appears
to reflect this pattern.
In summer, avoidance of the pipeline corridor
was primarily by cows and calves. Cows are most
sensitive to unusual stimuli just before par-
turition or during the early stages of labor (Lent
1966), but the present results suggest that
heightened sensitivity extends through the first 2
or 3 months post-partum. Other reports indicate
CAMERON ET AL.: CARIBOU DISTRIBUTION NEAR PIPELINE 16]
that cows and young calves are more easily
alarmed by, and more likely to flee from, a
potential threat than are male Caribou, barren
cows, or cows with older calves (de Vos 1960:
Lent 1966; Bergerud 1974). Loud noises, unless
associated with moving objects, do not generally
alarm Caribou which perceive and identify
adverse stimuli through visual and olfactory
means (Bergerud 1974). Cows with young calves,
however, consistently take flight from distant
stimuli, using visual analysis without verifica-
tion by scent (Bergerud 1974). Bergerud (1974)
and Lent (1966) reported that alert posture
assumed by individuals within a group did not
generally induce group flight unless cows and
neo-natal calves were present. Also, we noted
that cow-calf pairs on the calving grounds were
difficult to approach on foot to within 400 m.
Because of the importance of visual stimuh,
local terrain relief may influence the degree
of avoidance of the pipeline corridor. North
of Pump Station 2 the haul raod traverses
more than 100 km of continuous coastal plain
(see Figure 1) which is preferred summer range
for the Central Arctic herd. Construction ac-
tivity 1s frequently visible for 20 km or more, and
the greater inherent sensitivity of cows and
calves together with greater visibility provided
by flat terrain may account for the observed
avoidance of the haul road during summer. Bulls
do not appear to be as sensitive to local human
activity; however, since the mean latitude of
groups without calves (1.e., principally bulls) was
also displaced southward some evidence exists
for low-level avoidance of northern areas.
Avoidance of the Prudhoe Bay complex
continued during fall, and most Caribou were
observed farther inland. In September and
October groups along the haul road were well
mixed sexually, and cow-calf avoidance of the
pipeline corridor appears to have diminished.
This may be partially a result of a change in
group dominance associated with breeding;
rutting bulls clearly influence group activity
(Bergerud 1973; Cameron and Whitten, un-
published observations; Roby 1978) and may
alter the “normal” avoidance behavior of cow-
calf pairs. Perhaps more importantly, the ad-
vanced age of calves may have reduced the
sensitivity of maternal cows to human dis-
turbance.
162 THE CANADIAN FIELD-NATURALIST
Continued study is necessary to assess the
long-term responses of Caribou to oil develop-
ment in the Arctic. Sustained exposure of
Caribou to these stimuli may increase the degree
of avoidance, stabilize it at the current level, or
ultimately result in a reversal with subsequent
accommodation. Further, it is not known
whether this avoidance response, by virtue of its
probable influence on free movement, will
seriously alter herd productivity. Such are
matters for conjecture, and should be addressed
by future investigations.
Acknowledgments
This study was funded jointly by the Alaska
Department of Fish and Game through the Joint
State/ Federal Fish and Wildlife Advisory Team
and by Federal Aid in Wildlife Restoration
Project W-17-8. Logistic support by Alyeska
Pipeline Service Company is gratefully
acknowledged.
Literature Cited
Bergerud, A. T. 1973. Movement and rutting behavior of
caribou (Rangifer tarandus) at Mount Albert, Quebec.
Canadian Field-Naturalist 87: 357-369.
Bergerud, A. T. 1974. The role of the environment in the
aggregation, movement, and disturbance behavior of
caribou. Jn The behaviour of ungulates and its relation
to management. Volume 2. Edited by V. Geist and F.
Walther. International Union for Conservation of Nature
and Natural Resources Publications, New Series Number
24.
Vol. 93
Calef, G. W. 1974. The predicted effect of the Canadian
Arctic Gas pipeline project on the Porcupine caribou
herd. Jn Research reports, Volume IV, Environmental
impact assessment of the portion of the Mackenzie gas
pipeline from Alaska to Alberta. Environmental Protec-
tion Board, Winnipeg, Manitoba. pp. 101-120.
Cameron, R. D. and K. R. Whitten. 1979. Seasonal move-
ments and sexual segregation of caribou determined by
aerial survey. Journal of Wildlife Management (in press).
De Vos, A. 1960. Behavior of barren-ground caribou on
their calving grounds. Journal of Wildlife Management
24: 250-258.
Hemming, J. E. 1971. The distribution and movement
patterns of caribou in Alaska. Alaska Department of
Fish and Game, Wildlife Technical Bulletin Number 1.
60 pp. :
Klein, D. R. 1971. Reaction of reindeer to obstructions and
disturbances. Science 173: 393-398.
Lent, P. C. 1966. Calving and related social behavior in
the barren-ground caribou. Zeitschrift fur Tierpsycholo-
gie 6: 701-756.
Roby, D. D. 1978. Behavioral patterns of barren-ground
caribou of the Central Arctic herd adjacent to the Trans-
Alaska oil pipeline. M.Sc. thesis, University of Alaska.
200 pp.
Skoog, R. O. 1968. Ecology of caribou ( Rangifer tarandus
granti) in Alaska. Ph.D. thesis, University of California,
Berkeley, California. 699 pp.
White, R. G., B. R. Thomson, T. Skogland, S. J. Person,
D.E. Russell, D. F. Holleman, and J. R. Luick. 1975.
Ecology of caribou at Prudhoe Bay, Alaska. In Eco-
logical investigations of the tundra biome in the Prudhoe
Bay region, Alaska. Edited by J. Brown. Biological
Papers of the University of Alaska, Special Report
Number 2. pp. 150-201.
Received 4 October 1977
Accepted 27 December 1978
Interesting Vascular Plants from Southeastern
Yukon Territory
GEORGE W. SCOTTER! and WILLIAM J. CODY?
'Canadian Wildlife Service, Room 1000, 9942 — 108 Street, Edmonton, Alberta T5K 2J5
2Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario KIA 0C6
Scotter, George W. and William J. Cody. 1979. Interesting vascular plants from southeastern Yukon Territory. Canadian
Field-Naturalist 93(2): 163-170.
Recent vascular plant collections in extreme southeastern Yukon have added considerable data on the flora of the Yukon
Territory. This paper reports 19 taxa as newto the Yukon (Borrychium virginianum var. europaeum, Dryopteris spinulosa,
Matteuccia struthiopteris var. pensylvanica, Agrostis exarata, Cinna latifolia, Muhlenbergia mexicana, Oryzopsis
asperifolia, Poa ammophila, Sphenopholis intermedia, Parietaria pensylvanica, Vicia americana, Viola rugulosa, Circaea
alpina, Aralia nudicaulis, Osmorhiza depauperata, Castilleja miniata, Anaphalis margaritacea var. subalpina, Aster
ciliolatus, and Bidens cernua) and 43 taxa as extensions of known ranges into the southeastern Yukon.
Key Words: vascular plants, Yukon Territory, flora, range extensions.
During July of 1977, the senior author, in
connection with other work, had the oppor-
tunity to collect vascular plants from three sites
in extreme southeastern Yukon. Although 214
taxa of vascular plants were collected, the
purpose of this paper is to report only new
records and major extensions of known ranges
for the Yukon Territory.
Field studies were made at one cool spring and
two hot springs. The cool spring site, the most
intensively collected, was near two terraced tufa
deposits at the base of a limestone ridge on the
east side of the Coal River. Scalloped terrace-
faces on the west side of the pools were covered
with a variety of bryophytes and vascular plants
(Figure 1). Specimens from poorly drained
communities below the two tufa deposits and in
surrounding Black Spruce (Picea mariana),
White Spruce (P. glauca), Lodgepole Pine (Pinus
contorta var. latifolia), and Balsam Poplar
(Populus balsamifera) stands were also col-
lected.
The locations (Figure 2) and elevation of the
collection sites, date of collection, collectors’
names and collection numbers follow:
1. Cool springs and adjacent areas near the
Coal River, 60°09’N, 127°26’W, elevation
650-750 m, 16-19 July 1977, Scotter and T.
Ahti 24304-24598.
2. Hot springs and adjacent areas near Larsen
Creek, 60°12’N, 125°32’W, elevation 760 m,
20-22 July 1977, Scotter 24599-24756.
3. Hot springs and adjacent areas near the
Beaver River, 60°23’N, 125°34’W, elevation
450 m, 22 July 1977, Scotter 24757-25775.
The southeastern Yukon is mostly forested
with open stands of Black Spruce, White Spruce,
and Lodgepole Pine. Small portions of the area
extend above treeline, which occurs at approxi-
mately 1200-1350 m asl. The area lies within the
Liard and Hyland plateaus and the whole area
was covered by at least one advance of Cordil-
leran ice. For greater details on vegetation and
geology of the region readers are referred to
Douglas and Norris (1959), Bostock (1948),
Gabrielse and Blusson (1969), Rowe (1972), and
Oswald and Senyk (1977).
Voucher specimens have been deposited at the
Agriculture Canada herbarium (DAO). Unless
otherwise indicated, all specimens were deter-.
mined by Cody. A complete species list of the
plants collected is available and may be obtained
by contacting the authors.
In the annotated list, voucher numbers follow
the species; site numbers are in brackets. Species
not previously reported from the Yukon are
noted with an asterisk (*). Nomenclature follows
Porsild and Cody (1979).
The ecology of springs in northwestern Cana-
da has been little studied except for Liard Hot
Springs, British Columbia (Porsild and Crum
1961) and some near Hole-in-the-Wall Lake,
Nahanni National Park, Northwest Territories
(Arnold 1961). The three springs studied in the
Yukon are very different from each other in
structure and associated vegetation so that
163
Vol. 93
THE CANADIAN FIELD-NATURALIST
164
istic «tg land ali
Phe
AMOR SRS dal
Are
et
”
ia)
FIGURE |. The most common plants at the cool spring site growing on top of the tufa terraces were tall White Bog Orchid
(Hahenaria dilatata), Marsh Grass-of-Parnassus ( Parnassia palustris var. neogaea), Meadow Fireweed (Epi/obium
glandulosum var. adencaulon), Yellow Monkey Flower (Minulus guttatus), Pink Fleabane ( Erigeron philadelphicus),
Yellow Ragwort (Senecio pauperculus). and several species of grasses and sedges. Mats of velvety-green bryophytes
covered the sides of the moist tufa terraces.
1979
kilometres
Yukon
Territory
RR eS
Coal
River springs
SCOTTER AND CODY: VASCULAR PLANTS, YUKON TERRITORY 165
Nahanni National
Northwest
Territory
SAGAS ai) PRA
r
(
Beaver River \
springs
Larsen Creek \
springs
FIGURE 2. Map of the southeastern Yukon and adjacent regions showing the three collection sites in relation to Liard Hot
Springs and Nahanni National Park.
generalization on ecology is difficult. The Liard
Hot Springs and the springs near the Coal River.
however, have a close affinity in both structure
and associated vegetation.
*Botrychium virginianum var. europaeum (Rattle-
snake Fern). 24475, 24478 (1); 24712, 24673 (2).
Previously known from northern British Columbia
(Hultén 1968; Porsild 1951) and several sites in
southwestern District of Mackenzie (Cody 1963:
Scotter and Cody 1974).
Athyrium filix-femina ssp. cyclosorum (Lady Fern).
24473 (1).
This appears to be the second report of this species
from the Yukon Territory. Porsild (1951) found A.
filix-femina ssp. cyclosorum at a hot spring site in the
Mayo District. In the southwestern District of
Mackenzie it is known from only three localities
(Jeffrey 1961; Porsild 1961; Scotter and Cody 1974),
two of which are hot spring sites.
*Drvopteris spinulosa
24468, 24472, 24482 (1).
(Spinulose Wood Fern).
166
Rare in the northwestern part of its range. A site in
Nahanni National Park, the second site in the District
of Mackenzie, is soon to be reported by Cody et al.’
* Matteuccia struthiopteris var. pensylvanica (Pteritis
nodulosa) (Ostrich Fern). 24764 (3).
According to Hultén (1968), this fern is restricted to
small isolated areas in Alaska. In northern British
Columbia, Porsild and Crum (1961) reported that it
formed dense colonies near and below the springs at
Liard Hot Springs. In the southwestern District of
Mackenzie it occurs on the flood plain of the Liard
River nearly as far downstream as Nahanni Butte
(Cody 1963) and onthe floodplains of the lower South
Nahanni River (Scotter and Cody 1974).
Equisetum hyemale var. affine (Scouring Rush).
24743 (2).
According to the map in Hultén (1968), this
collection extends the known range to the south-
eastern Yukon Territory.
Selaginella selaginoides (Spikemoss). 24527 (1); 24639
(2).
These collections of this species in the southeastern
Yukon Territory change the position of the delimiting
line in Hultén (1968) to include all of the southern
Yukon Territory.
x Agrohordeum macounti (Elymus macounii). 24617
(2).
This hybrid of Agropyron trachycaulum and
Hordeum jubatum was known inthe Yukon Territory
by Porsild (1951) from only three collections. In the
herbarium of the Department of Agriculture at
Ottawa (DAO), there are now an additional eight
collections. The hybrid is thus of much more frequent
occurrence than previously suspected, and may occur
wherever the two parents are found growing in close
proximity.
* Agrostis exarata. 24345, 24351, 24358 (1); 24622B
(2).
Cody and Porsild (1968) extended the known range
of this species from the Pacific Coast to the District of
Mackenzie at Tungsten. Scotter and Cody (1974) have
since reported additional sites at both hot and cool
springs in Nahanni National Park.
Bromus ciliatus (Fringed Brome). 24320, 24332 (1);
24610, 24611 (2).
Hultén (1968) shows only three widely separated
Yukon sites on his map. The species is not common in
3Cody, W. J., G. W. Scotter, and S. S. Talbot. Additions
to the vascular plant flora of Nahanni National Park,
Northwest Territories (in preparation).
THE CANADIAN FIELD-NATURALIST
Vol. 93
southwestern District of Mackenzie, but Porsild
(Porsild and Crum 1961) has collected it at Liard Hot
Springs.
* Cinna latifolia (Wood Reedgrass). 24412 (1).
Cody (1963) reported a number of stations along
the lower Liard River, and Porsild and Crum (1961)
found it to be occasional in rich mixed aspen and
spruce woods at Liard Hot Springs.
Danthonia intermedia (Wild Oat-Grass). 24770 (3).
This collection helps fill in the known distribution
of this species between sites in southern Yukon
Territory (Porsild 1951, 1966) shown on Hultén’s
(1968) map, and sites reported by Scotter and Cody
(1974) and Cody et al.3 in Nahanni National Park.
Glyceria striata var. stricta (Fowl Manna Grass).
24342, 24416, 24523, 24555 (1); 24625 (2).
Porsild (1951) surmised that this grass would be
found in the Yukon Territory because !t was already
known from central Alaska and the Liard Hot
Springs. Hultén (1968) depicted a collection from near
Watson Lake (Calder & Kukkonen 27617 and Calder
& Gillett 26544 [DAO]). The specimens cited above
thus represent the second report of this species for the
Yukon Territory.
* Muhlenbergia mexicana. 24629 (2). (det. C.G.
Reeder).
Porsild (Porsild and Crum 1961) and Welsh (1974)
reported this grass from the Liard Hot Springs.
* Oryzopsis asperifolia (Mountain-Rice). 24435 (1).
In northern British Columbia, O. asperifolia is
known from Mile 578 Alaska Highway, 59°38’N,
127°29’W (Calder & Gillett 25653 [DAO]). In the
District of Mackenzie it has recently been reported
from Nahann: National Park (Cody et al.3), an
extension of range to the northwest from the Salt
Plains west of Fort Smith (Raup 1936).
Phalaris arundinacea (Reed Canary Grass). 24408 (1).
The map in Hultén (1968) depicts collections from
Whitehorse (Porsild 1951) and near Teslin Lake.
Additional Yukon Territory specimens at DAO are
from Albert Creek, Mile 643.5 Alaska Highway
(Calder & Gillett 26545, Harms 17199).
* Poa ammophila. 24596 (1). (det. M. Barkworth).
This is an extension of the known range of some
900 km to the south from the type region at the mouth
of the Mackenzie River on the arctic coast of the
District of Mackenzie. It 1s not known to occur
between those collections.
Schizachne purpurascens. 24434 (1).
Previous collections in the Yukon Territory were
from along the Canol Road (Porsild 1951; Hultén
1968).
1979
* Sphenopholis intermedia. 24314 (1); 24602, 24612,
24740 (2).
Porsild (1951) and Hultén (1968) reported this grass
from Liard Hot Springs and Tanana (Manley) Hot
Springs in Alaska. Scotter and Cody (1974) reported
it from Nahanni National Park and commented on its
presence elsewhere in the District of Mackenzie.
Carex atherodes. 24510, 24511 (1).
These collections will move the delimiting line on
the map in Hultén (1968) to the east to include the
extreme southeastern section of the Yukon Territory.
Carex concinna. 24452 (1).
This collection will move the delimiting line on the
map in Hultén (1968) to the east to include the
extreme southeastern section of the Yukon Territory.
Juncus balticus var. littoralis (J. arcticus spp. ater,
sensu Hultén 1968) (Wire Rush). 24540 (1).
This collection will move the delimiting line on the
map in Hultén (1968) to the east to include the
extreme southeastern section of the Yukon Territory.
Allium schoenoprasum var. sibiricum (Wild Chives).
24563 (1).
This collection will move the delimiting line on the
map in Hultén (1968) to the east to include the
extreme southeastern section of the Yukon Territory.
Smilacina trifolia (Three-leaved Solomon’s-seal).
MTS 3) (De
Apparently rare in southeastern Yukon Territory.
The single dot onthe map in Hultén (1968) is probably
based on specimens gathered at Watson Lake by
Raup and Correll (Porsild 1974). It was also collected
there by Mitchell 9] and Gillett 3618 (DAO). Another
unreported collection is from the Cassiar Mountains,
60°18’N, 130°18’W, Poole 27 (DAO).
Goodyera repens (Rattlesnake Plantain). 24437 (1).
According to the map in Hultén (1968), previous
collections from the Yukon Territory have ali been
made west of 130° W longitude.
Salix longistylis (S. alaxensis var. longisiylis). 24584
(1): 24611, 24644, 24662 (2).
These collections will move the delimiting line on
the map in Hultén (1968) to the east to include the
extreme southeastern section of the Yukon Territory.
The species is, as well, based on collections at DAO,
much more frequent in the Mackenzie Mountains in
the western District of Mackenzie, than indicated on
that map.
Salix padophylla. 24518, 24576 (1); 24660 (2).
These collections will move the delimiting line on
the map in Hultén (1968) eastwards to include the
extreme southeastern section of the Yukon Territory.
To the east it has been recorded from several sites in
Nahanni National Park (Scotter and Cody 1974).
SCOTTER AND CODY: VASCULAR PLANTS, YUKON TERRITORY 167
* Parietaria pensylvanica (Pellitory). 24725 (2).
The distribution of this species in Canada is quite
spotty (Bassett et al. 1974) because of its apparently
restricted habitat requirement of cool, moist, shady,
rocky situations. Porsild (1961) reported the ex-
tension of the known range from central British
Columbia north to Liard Hot Springs, and discussed
the distribution.
Urtica gracilis (Nettle). 24462 (1); 24732 (2).
Apparently rare in the Yukon Territory. Hultén
(1968) indicates collections from the Dawson area,
and near the British Columbia border.
Polygonum amphibium var. stipulaceum (Water
Smartweed). 24413, 24513 (1).
These collections will move the delimiting line on
the map in Hultén (1968) eastwards to include the
extreme southeastern section of the Yukon Territory.
Minuartia dawsonensis. 24365, 24372, 24592B (1);
24604, 24641 (2).
These collections will move the delimiting line on
the map in Hultén (1968) eastwards to include the
extreme southeastern part of the Yukon Territory. It
is also known from Nahanni National Park (Scotter
and Cody 1974).
Silene menziesii. 24483 (1).
On the basis of the distribution map in Hultén
(1968), this species appears to have a broken
distribution in the northern part of its range. This
collection will move the delimiting line on the map in
Hultén (1968) to the east to include the extreme
southeastern part of the Yukon Territory.
Ranunculus abortivus (Kidneyleaf Buttercup). 24605
(2).
This is apparently a rare species in the Yukon
Territory. Hultén (1968) shows only four dots from
the south central part of the Territory.
Thalictrum sparsiflorum var. richardsonii (Meadow-
Rue). 24406, 24551 (1).
These collections, from the extreme southeastern
part of the Yukon Territory, are intermediate between
the sites on the map in Hultén (1968) and the Liard
River sites in southwestern District of Mackenzie
reported by Jeffrey (1961).
Amelanchier alnifolia (Saskatoon). 24422, 24594 (1);
24682 (2).
These collections will move the delimiting line on
the map in Hultén (1968) eastwards to include the
extreme southeastern part of the Yukon Territory.
Dryas drummondii (Yellow Dryad). 24590 (1): 24626
(2).
The collections, from the extreme southeastern part
of the Yukon Territory, are from an area inter-
168
mediate between two areas of distribution outlined by
Hultén (1968).
Geum aleppicum var. strictum (Avens). 24618, 24731
(2).
Hultén (1968) shows only a single disjunct locality
in southwestern Yukon Territory. The main range
extends across Canada from the east coast and
reaches Nahanni National Park in southwestern
District of Mackenzie.
Sanguisorba sitchensis (S. stipulata, sensu Hultén
(1968)) (Burnet). 24415 (1).
This collection, from the extreme southeastern part
of the Yukon Territory, will move the delimiting line
on the map in Hultén (1968) to the east and thus
include all of the southern Yukon.
Astragalus americanus. 24564 (1).
According to Hultén (1968), previous collections
from the Yukon Territory have all come from west of
longitude 132°W.
Oxytropis deflexa var. foliolosa. 24646 (2).
This collection helps complete our knowledge of the
distribution of var. foliolosa in the Yukon Territory
by extending the delimiting line around the southern
population to the extreme southeastern Yukon. The
map in Hultén (1968) depicts widely separated
northern and southern populations in the Yukon
Territory.
*Vicia americana (Wild Vetch). 24745 (2).
This species occurs as far north in British Columbia
as Liard Hot Springs (Porsild and Crum 1961) while
in southwestern District of Mackenzie it is found
along the Liard River (Jeffrey 1961) and in Nahanni
National Park (Scotter and Cody 1974).
Geranium richardsonti. 24747 (2).
Hultén (1968) shows only a few collection sites in
central Yukon Territory and the hot springs at
Tungsten in the headwaters of the Flat River in the
District of Mackenzie. It is also known from Hole-in-
the-Wall Lake in Nahanni National Park (Porsild
1961) in the District of Mackenzie.
Viola langsdorffti. 24638, 24726 (2).
Hultén (1968) indicates two inland sites in central
and southern Yukon Territory which are remote from
the mainly coastal range of the species. The collec-
tions cited here represent another site more remote
from the main range.
*Viola rugulosa (Western Canada Violet). 24735 (2).
This is a westward extension of range from sites in
the Liard and South Nahanni valleys in southwestern
District of Mackenzie reported by Jeffrey (1961) and
Scotter and Cody (1974).
THE CANADIAN FIELD-NATURALIST
Vol. 93
*Circaea alpina (Enchanter’s Nightshade). 24758 (3).
This circumpolar species is known from northern
British Columbia at Liard Hot Springs (Porsild and
Crum 1961), and in the adjacent District of Mac-
kenzie in the headwaters of the Flat River and along
the Liard River (Scotter and Cody 1974).
* Aralia nudicaulis (Wild Sarsaparilla). 24427 (1).
Porsild and Crum (1961) found this species at Liard
Hot Springs, while Jeffrey (1961) and Scotter and
Cody (1974) reported it from several sites in the
southwestern District of Mackenzie.
Cicuta maculata var. angustifolia (Water Hemlock).
24739 (2).
Only one other collection is known from the Yukon
Territory by the authors: Tatchun River, Mile 117.5
Mayo Highway, Taylor 4080 (DAO).
* Osmorhiza depauperata (O. obtusa) (Sweet Cicely).
24479 (1); 24687 (2).
Recorded from adjacent northern British Columbia
at Liard Hot Springs by Porsild and Crum (1961), and
from the Liard River Valley in southwestern District
of Mackenzie by Cody (1963).
Collomia linearis. 24730, 24742 (2).
This is a native North American species which
Hultén (1968) considers to be introduced inthe Yukon
Territory. It seems doubtful that it was introduced to
this remote and pristine site. Previous collections,
according to Hultén’s map, are all west of longitude
135°W.
Scutellaria galericulata var. pubescens (Skullcap).
2ATTLA (3).
Previously known from the Yukon Territory only
from the vicinity of Mayo (Porsild 1951, 1974; Hultén
1968).
Castilleja caudata. 2456S (1).
This collection extends the known range of this
species to the southeast from western and northern
Yukon Territory.
* Castilleja miniata (Red Indian Paint-brush). 24683,
24728 (2).
This is a northward extension of range from north
central British Columbia.
Mimulus guttatus (Yellow Monkey-flower). 24306,
24339 (1); 24678 (2).
According to Hultén (1968) and Porsild (1974) this
species barely enters the southern part of the Yukon
Territory near Whitehorse and Watson Lake. It has
been reported from mineral springs in Nahanni
National Park (Scotter and Cody 1974). The plant
was very abundant at the Coal River springs.
1979
Galium triflorum (Sweet-scented Bedstraw). 24516
(1); 24675 (2).
This species, according to the map in Hultén (1968),
barely enters the southern part of the Yukon Territory
west of 133°W longitude. Our collections will move
the delimiting line eastward to include the extreme
southeastern Yukon Territory. In the adjacent Dis-
trict of Mackenzie it is known along the Liard River
(Cody 1961: Jeffrey 1961) and in Nahanni National
Park (Scotter and Cody 1974); in northern British
Columbia it has been found at Liard Hot Springs
(Porsild and Crum 1961).
* Anaphalis margaritacea var. subalpina (Pearly Ever-
lasting). 24727 (2).
The site at Hole-in-the-Wall Lake, Nahanni Na-
tional Park, reported by Porsild (1961) was widely
disjunct from the Alaska Panhandle sites shown on
the map in Hultén (1968). The specimen reported here
helps fill the gap between these widely separated areas.
Arnica cordifolia. 24387 (1).
This collection comes from an area intermediate
between the sites shown on the map in Hultén (1968),
and an apparently isolated site on the lower Liard
River reported by Raup (1947).
Aster alpinus ssp. vierhapperi. 24783 (1).
This collection from the extreme southeastern part
of the Yukon Territory will move the delimiting line in
Hultén (1968) to the east, thus bringing it close to the
populations in southwestern District of Mackenzie
cited by Scotter and Cody (1974).
* Aster ciliolatus. 24738 (2): 24775 (3).
These collections extend the known distribution of
A. ciliolatus to the west from Fort Liard on the Liard
River in southwestern District of Mackenzie (Jeffrey
1961).
Aster modestus. 24399 (1): 24356, 24648, 24784 (2).
Hultén (1968) indicated a collection from near
Dawson and Porsild and Crum (1961) reported this
aster at Liard Hot Springs in northern British
Columbia.
* Bidens cernua (Nodding Beggar-ticks). 24767, 24774
(3).
In central Alaska this species is known from the
Yukon River drainage (Hultén 1968) while in the
District of Mackenzie it is known from Nahanni
National Park, Great Slave Lake, and the Slave River.
Erigeron philadelphicus (Pink Fleabane). 24336,
24340, 24350 (1); 24608, 24647, 24676 (2).
Hultén (1968) indicates a single Yukon Territory
locality near Old Crow, from whence it has also been
reported by Wein et al. (1974). Porsild and Crum
(1961) reported it at Liard Hot Springs.
SCOTTER AND CODY: VASCULAR PLANTS, YUKON TERRITORY 169
Hieracium scabriusculum. 24757, 24759B (3).
Hultén (1968) indicates two sites for this species,
one near Watson Lake and another near Dawson.
Solidago canadensis var. salebrosa (Canadian Gol-
denrod). 2474] (2).
Hultén (1968) indicates collections from the vicinity
of Dawson and Watson Lake.
Literature Cited
Arnold, E. W. 1961. Plant communities of a hot spring
in the Mackenzie Mountains, Northwest Territories.
M.Sc. thesis, University of Michigan, Ann Arbor. 34 pp.
Bassett, I. J.. C. W. Crompton, and D. W. Woodland. 1974.
The family Urticaceae in Canada. Canadian Journal of
Botany 52: 503-516.
Bostock, H.S. 1948. Physiography of the Canadian Cor-
dillera, with special reference to the area north of the
fifty-fifth parallel. Geological Survey Canada, Memoir
247. 106 pp.
Cody, W. J. 1961. New plant records from the upper
Mackenzie River Valley, Mackenzie District, Northwest
Territories. Canadian Field-Naturalist 75: 55-69.
Cody, W. J. 1963. A contribution to the knowledge of the
flora of southwestern Mackenzie District. Canadian
Field-Naturalist 77: 108-123.
Cody, W. J.and A. E. Porsild. 1968. Additions to the flora
of continental Northwest Territories, Canada. Canadian
Field-Naturalist 82: 263-275.
Douglas, R. J.W. and D. K. Norris. 1959. Fort Liard and
La Biche map-areas, Northwest Territories and Yukon.
Geological Survey Canada, Paper 59-6. 23 pp.
Gabrielse, H. and S. L. Blusson. 1969. Geology of Coal
River map-area, Yukon Territory and District of Mac-
kenzie. Geological Survey Canada, Paper 68-38. 22 pp.
Hultén, E. 1968. Flora of Alaska and neighboring terri-
tories. Stanford University Press, Stanford. 1008 pp.
Jeffrey, W. W. 1961. Notes on plant occurrence along
lower Liard River, Northwest Territories. National Mu-
seum of Canada Bulletin 171: 32-115.
Oswald, E. T. and J. P. Senyk. 1977. Ecoregions of Yukon
Territory. Department of Fisheries and Environment, —
Canadian Forestry Service Report BC-X-164. 115 pp.
Porsild, A. E. 1951. Botany of southeastern Yukon adja-
cent to the Canol Road. National Museum of Canada
Bulletin 121: 1-400.
Porsild, A. E. 1961. The vascular flora of an alpine valley
in the Mackenzie Mountains. National Museum of
Canada Bulletin 171: 116-130.
Porsild, A. E. 1966. Contributions to the flora of south-
western Yukon Territory. National Museum of Canada
Bulletin 216: 1-86.
Porsild, A. E. 1974. Materials for a flora of central Yukon
Territory. National Museum of Canada, Publications in
Botany, Number 4: 1-77.
Porsild, A. E. and H. Crum. 1961. The vascular flora of
Liard Hot Springs, B.C., with notes on some bryophytes.
National Museum of Canada Bulletin 171: 131-197.
Porsild, A. E. and W. J. Cody. 1979. Vascular plants of
continental Northwest Territories. National Museums of
Canada, Publications in Botany, Number 8. (in press.)
170 THE CANADIAN FIELD-NATURALIST
Raup, H. M. 1936. Phytogeographic studies in the Atha-
baska-Great Slave Lake region. I, Catalogue of the
vascular plants. Journal of the Arnold Arboretum 17:
180-315.
Raup, H.M. 1947. The botany of southwestern Mac-
kenzie. Sargentia 6: 1-275.
Rowe, J. S. 1972. Forest regions of Canada. Department
of the Environment, Canadian Forest Service Publication
Number 1300. 172 pp.
Scotter, G. W. and W. J. Cody. 1974. Vascular plants of
Nahanni National Park and vicinity, Northwest Terri-
tories. Naturaliste Canadien 101: 861-891.
Vol. 93
Wein, R. W., L. R. Hettinger, A. J. Janz, and W. J. Cody.
1974. Vascular plant range extensions in the northern
Yukon Territory and northwestern Mackenzie District.
Canada. Canadian Field-Naturalist 88: 57-66.
Welsh, S. L. 1974. Anderson’s flora of Alaska and adjacent
parts of Canada. Brigham Young University Press, Provo,
Utah. 724 pp.
Received 15 August 1978
Accepted 22 January 1979
Notes
Fall Foods of Common Snipe on the Copper River Delta, Alaska
STANLEY E. SENNER and P. G. MICKELSON
Institute of Arctic Biology, University of Alaska, Fairbanks, Alaska 99701
Senner, Stanley E. and P. G. Mickelson. 1979. Fall foods of Common Snipe on the Copper River Delta, Alaska. Canadian
Field-Naturalist 93(2): 171-172.
Stomachs from 22 Common Snipe (Capella gallinago delicata) collected at three sites on the Copper River Delta, Alaska,
were examined to determine food habits. Cranefly larvae, Prionocera sp.(1 ipulidae) were the most important food items and
occurred in 75 to 86% of the stomachs. These and beetle larvae were the major foods, as previously shown for other breeding
and staging areas in Canada.
Key Words: Common Snipe, food habits, Copper River Delta, Alaska.
Common Snipe (Capella gallinago delicata) breed
primarily across boreal North America (Tuck 1972),
although some are found breeding in arctic regions
(e.g., Colville River, Alaska (Kessel and Cade 1958)).
Most literature on snipe in Alaska concerns their
distribution (e.g., Gabrielson and Lincoln 1959).
There are no previous studies on food habits of snipe
in Alaska, although Tuck (1972) described food habits
of snipe at breeding and fall concentration sites in
Canada. During fall the Copper River Delta in south-
central Alaska may host several 10 000s of snipe
(isleib and Kessel 1973). The majority of these migrate
south, presumably to wintering grounds from British
Columbia to California (Burleigh 1955; Tuck 1972),
although snipe rarely have been recorded in mid-
winter on the western Copper River Delta (Isleib and
Kessel 1973).
Study Area
Snipe were collected at three sites on the Copper
River Delta. Nine snipe were from Mile 20 of the
Copper River Highway (60°27’N, 145°16’W). Carex
spp., Eleocharis spp., Equisetum spp., and Hippuris
vulgaris dominated this meadow habitat. Five snipe
were collected in mixed sedge-grass-forb habitat
dominated by Carex lyngbyaei, Deschampsia sp.,
Festuca rubra, and Hedysarum alpinum along lower
Alaganik Slough (60°26’N, 145°17’W). Eight snipe
were from the east Copper River Delta (60°17’N,
144°56’W) in habitat dominated by A/nus crispa,
Myrica gale, and Carex lyngbyaei.
Methods
All snipe were taken between | September and 10
October in 1976 (10 specimens) and 1977 (12
specimens). Contents of the snipe gizzard and
proventriculus, and sometimes the esophagus, were
removed and placed in Whirl-paks with hexamethy-
lenetetramine-buffered formalin (Mueller 1972).
Time between collection of the bird and removal of
stomach contents varied up to several hours. Because
of differential digestion rates and post-mortem
digestion (e.g., Tuck 1972), only the number of
stomachs with recognizable items are reported here.
Tuck (1972) suggested that frequency of occurrence
(Hartley 1948) might be more meaningful than
volume or weight for presenting food habits data. Our
results are expressed in frequency of occurrence
because volume or weight assigns great importance to
large or difficult-to-digest items of food and
underemphasizes the importance of small or easily
digested food items. Vegetable debris, seeds, and
mineral grit were not considered as food (Tuck 1972).
White and Harris (1966) and Tuck (1972) found that
plant fibers and seeds remained relatively unchanged
by any digestive process and were finally regurgitated.
Results and Discussion
Nineteen of 22 snipe collected on the Copper River
Delta contained food items (Table |). The most
important food items, represented by up to 43 |
individuals in a single stomach, were Cranefly larvae,
Prionocera sp. (Tipulidae), which were present in 75
to 86% of the stomachs from all three sites (Table 1).
The mean length of 26 intact larvae was
24.4 + 5.82 mm.
Beetles, especially larval Dytiscidae (probably
Agabus sp.) were the second most important item in
the diet of snipe during fall (Table 1). These occurred
in 14 to 75% of the stomachs from snipe collected from
the three sites. Both cestodes and nematodes were
found in snipe stomachs at Mile 20 (Copper River
Highway) and Alaganik Slough, but neither was
found in snipe taken on the east Copper River Delta.
The nematodes did not appear to be parasitic on the
snipe.
The results of this study agree with those sum-
marized by Tuck (1972) in that insects, particularly
171
172
TABLE 1—Frequency of occurrence of food items in the
stomachs of 19 Common Snipe collected at three sites on the
Copper River Delta, Alaska
Percent frequency of occurrence
East
Copper
Alaganik River
Food Mile 20 Slough Delta
categories (N=8) (N=4) (N=7)
Animal taxa
Cestoda IDES 50.0 —
Nematoda 25.0 25.0 —
Lumbricidae 12.5 -— —
Gastropoda, undetermined 12.5 _ —
Pelecypoda, undetermined 12.5 _ _—
Insecta, undetermined — — 14.3
Diptera (total) 75.0 100.0 85.7
Tipulidae
Prionocera sp. larvae 75.0 75.0 85.7
Pedicia sp. larvae a 25.0 —
Chironomidae larvae = 25.0 28.6
Coleoptera (total) 50.0 75.0 14.3
Undetermined adult — 25.0 14.3
Dytiscidae larvae
(largely Agabus sp.?
and Hydroponidae) 375 75.0 14.3
Carabidae adult 12.5 — ---
Staphylinidae adult — — 14.3
Araneida, undetermined a 25.0 —
Non-food items!
Vegetable debris 87.5 75.0 100.0
Seeds 87.5 75.0 85.7
Mineral grit 100.0 75.0 71.4
1See Tuck (1972).
Diptera and Coleoptera larvae, are taken frequently.
Tuck (1972) mentions that Tipulidae are especially
significant, primarily on snipe breeding grounds, but
also on fall staging areas in Canada. In northern
latitudes Tipulidae are of great importance to other
scolopacid shorebirds (see Chernov 1967; Holmes and
Pitelka 1968). Our study has confirmed their value on
THE CANADIAN FIELD-NATURALIST
Vol. 93
an important staging and breeding ground for
Common Snipe along the north Gulf of Alaska coast.
Acknowledgments
We thank Daniel D. Roby, William A. Lehn-
hausen, John W. Smith, Brian E. Lawhead, and
Stephen Murphy for assistance in collecting snipe.
George J. Mueller, Stephen F. MacLean, Jr., and
Jacqueline D. LaPerriére helped identify invertebrate
specimens. Stephen F. MacLean, Jr., James C.
Bartonek, Leslie Tuck, and D. Keppie reviewed the
manuscript.
Literature Cited
Burleigh, T. D. 1955. Investigations of woodcock, snipe and
rails in 1954. U.S. Fish and Wildlife Service Special
Scientific Report, Wildlife 28. 62 pp.
Chernov, Y.I. 1967. [Trophic relationships of birds to
insects in the tundra zone.] Ornitologi’ya 8: 133-150.
(Translated from Russian by S. F. Maclean, Jr.)
Gabrielson, I. N. and F. C. Lincoln. 1959. Birds of Alaska.
Wildlife Management Institute, Washington, D.C. 922 pp.
Hartley, P. H. 1948. The assessment of the food of birds.
Ibis 90: 361-381.
Holmes, R.T. and F.A. Pitelka. 1968. Food overlap
among coexisting sandpipers on northern Alaskan tundra.
Systematic Zoology 17: 305-318.
Isleib, M. E. “Pete” and B. Kessel. 1973. Birds of the North
Gulf Coast-Prince William Sound Region, Alaska.
Biological Papers of the University of Alaska Number 14.
149 pp.
Kessel, B. and T. J. Cade. 1953. Birds of the Colville River,
northern Alaska. Biological Papers of the University of
Alaska Number 2. 83 pp.
Mueller, G. J. 1972. Field preparation of marine specimens.
University of Alaska Museum, Fairbanks. 44 pp.
Tuck, L. M. 1972. The snipes. Canadian Wildlife Service
Monograph Number 5. 428 pp.
White, M. and S. W. Harris. 1966. Winter occurrence,
foods, and habitat use of snipe in northwest California.
Journal of Wildlife Management 30(1): 23-34.
Received 5 April 1978
Accepted 23 November 1978
1979
NOTES 73}
Ivory Gull Colonies in Southeastern Ellesmere Island, Arctic Canada
T. FRISCH and W. C. MORGAN
Geological Survey of Canada, 588 Booth Street, Ottawa, Ontario KIA 0E4
Frisch, T. and W. C. Morgan. 1979. Ivory Gull colonies in southeastern Ellesmere Island, Arctic Canada. Canadian
Field-Naturalist 93(2): 173-174.
Five Ivory Gull colonies, ranging in population from about 12 to 60 birds, have been discovered in the upland icefields of
southeastern Ellesmere Island. The colonies occur on cliff-faces of precipitous nunataks as much as 26 km from the nearest
sea coast. One colony is established as a nesting site; the others are probable breeding places.
Key Words: Ivory Gull, Ellesmere Island, Arctic, colonies, nunatak nesting site.
Only one active breeding colony of the Ivory Gull
(Pagophila eburnea) has heretofore been known from
the North American Arctic: Seymour Island, a small
low island north of Bathurst Island (MacDonald
1976). In the summer of 1977, while engaged in a
helicopter-supported geological survey, the authors
found five colonies of Ivory Gulls in the highlands of
southeastern Ellesmere Island, an environment very
different from that of Seymour Island. This note
records the newly-discovered localities and our
observations.
The area of Ellesmere Island that contains the
colonies is bounded by latitude 78°15’N and longi-
tude 81° W and traversed by a major fiord, Makinson
Inlet (Figure 1). Most of the land area comprises
upland icefields and has a maximum relief of 1500 m;
rock exposures are found mainly in coastal cliffs and
nunataks. The terrain shown in Figure 2 lies north of
Makinson Inlet between colonies 4 and 5 (Figure 1)
and is representative of much of the area, which is
underlain largely by metamorphic and granitic rocks
of the Canadian Shield. The only permanent human
settlement in the area is the small Inuit community of
Grise Fiord, on the south coast of Ellesmere Island.
Positions of the five Ivory Gull colonies discovered
are plotted on Figure | and more accurate locations in
the Universal Transverse Mercator Grid system on
National Topographic System 1:250 000 maps are
given below. All sites are on rock ledges near the tops
of steep cliff-faces of nunataks several hundred metres
high at elevations of 450 to 750 m above sea-level.
Only site 2, which was located purely by chance after
the helicopter had landed and shut down, was
observed at relatively close range. Owing to their
inaccessibility, and lack of time and/or the threat of
excessive disturbance by the helicopter, the other sites
could not be examined closely. Numbers of gulls seen
are very approximate and are based chiefly on
estimates made during a single fly-past.
Information on each colony is listed in the
following order: NTS 1:250 000 map sheet; approxi-
mate location on map; UTM easting and northing in
Zone 17; approximate elevation above sea-level;
distance to nearest sea coast; date of sighting;
observer. Other details follow.
Site 1. Craig Harbour 49A; NE corner; 508600, 8532300;
750 m; 23 km; 11 August; TF. Sixty birds flying around cliff
ima “80 u TI ee
jee Bae ~~ =
q eS = Adogan Inlet
as Ss
Lagi ny!
BS tie
eC nN 184
4, Ne ee
ae Talbot Inlet SS } y
ww ‘eas
a
4
i
Ge
? BAF FIN
BaeAw
114
\,
(¢
YW
)Clarence
( Head
\
i
Bes) ee
5 ey
We = ¢
aS (
X ff ie i
pa ire
we
)
(
;
25km
i J 1 1
FIGURE |. Sketch-map of southeastern Ellesmere Island
showing five sites of Ivory Gullcolonies. Broken lines
mark the edges of icefields and glaciers but nunataks
are not shown.
174
THE CANADIAN FIELD-NATURALIST
Vol93
FIGURE 2. Typical terrain north of site 4.
and resting on ledges. A flat area on top of the nunatak was
free of snow but not occupied by birds.
Site 2. Clarence Head 39B; NW corner, 526000, 8537400;
450 m; 14km; 12 August; WCM. Twelve to 15 birds seen
near top of cliff. Four large young birds visible by eye in nests
on excrement-stained ledges. No unhatched eggs were visible
through binoculars. Observations were made from an
icefield looking down on the site across a gully.
Site 3. Talbot Inlet 39C and part of 39D; Thorndike Peaks;
541200, 8565000; 600 m; 6 km; 12 August; WCM. Fifty birds
in the air, resting on ledges, and taking off from ledges.
Site 4. Talbot Inlet 39C and part of 39D; Inglefield
Mountains; 543600, 8597600; 600 m; 12 km; 29 June; WCM.
At least 50 birds suddenly encountered when helicopter flew
around corner of nunatak.
Site 5. Ekblaw Glacier 39F and 39E; head of Cadogan
Glacier; 549900, 8679000; between 600 and 750 m; 26 km; 15
August; TF. Thirty birds on ledges and flying around
nunatak.
Site 2, southwest of Smith Bay (Figure 1), is thus
established as a new breeding place of the Ivory Gull.
Although we have no direct evidence of breeding at
the other localities, we consider it more than probable
that these are also nesting sites.
Nesting sites of the Ivory Gull similarly situated on
nunataks well inland from the coast have been
reported from southern Vestspitsbergen by Birken-
majer and Skreslet (1963) (see also Lovenskiold 1964).
No doubt sites of this type are used because, among
other factors, they provide a haven safe from
predators. Furthermore, Polar Bear (Ursus mari-
timus) kills provide a major source of food for the
Ivory Gull (MacDonald 1967, p. 6) and all the
Ellesmere Island sites are relatively near the east coast,
which, according to our observations, supports a
sizable bear population, particularly in Smith Bay.
Our chiefly aerial coverage of southeastern Elles-
mere Island was by no means complete and, judged
from essentially chance discoveries of five colonies
and a number of sightings of individual Ivory Gulls, it
seems certain that more colonies exist in the upland
icefields. Also, our sightings lend credence to uncon-
firmed reports from the 19th century. of a breeding
place on the east coast of Ellesmere Island, north of
the area described here (MacDonald and Macpherson
1962). In any event, southeastern Ellesmere Island is
now established as an important breeding ground of
the Ivory Gull.
Acknowledgments
We thank S. D. MacDonald, D. N. Nettleship, and
H. Ouellet for reviews of this note and the Polar
Continental Shelf Project for support in the field.
Literature Cited
Birkenmajer, K. and S. Skreslet. 1963. Breeding colony of
ivory gulls in Torell Land, Vestspitsbergen. Norsk Polar-
institutt Arbok 1962: 120-126.
Lovenskiold, H. L. 1964. Avifauna Svalbardensis. Norsk
Polarinstitutt Skrifter 129.
MacDonald, S. D. 1976. Phantoms of the polar pack ice.
Audubon 78(3): 2-19.
MacDonald, S. D. and A. H. Macpherson. 1962. Breeding
places of the Ivory Gull in Arctic Canada. National
Museum of Canada Bulletin 183. pp. 111-117.
Received 23 August 1978
Accepted 17 November 1978
79
NOTES 175
Wind-caused Death of Great Cormorant
MARTIN K. MCNICHOLL! and GEOFFREY G. HOGAN
Department of Biological Sciences, Brock University, St. Catharines, Ontario L2S 3A1
'Present Address: Beak Consultants Ltd., 3530 11 A St. N.E., Calgary, Alberta T2E 6M7
MeNicholl, Martin K. and Geoffrey G. Hogan. 1979. Wind-caused death of Great Cormorant. Canadian Field-
Naturalist 93(2): 175.
Key Words: Accidental death, Great Cormorant, Prince Edward Island, weather, wind.
At 18:32 on 10 April 1977, McNicholl watched an
adult Great Cormorant (Phalacrocorax carbo) fly
from the open sea to ledges near the top of a
cormorant colony cliff at Cape Tryon on the northern
shore of Prince Edward Island. Strong north winds,
rated as 5 or more on the Beaufort scale
( = 29-38+ km/h), were blowing at the time, and
visibility was reduced by blowing snow, although the
snow also heightened light levels. The bird tried three
times to land, but each time gusts caused it to circle
back towards the sea. On the fourth approach, the
bird was gliding into a position from which it could
land when a sudden gust of wind threw it breast-first
into the cliff face. The bird flapped one wing twice,
then ceased all motion. A Common Raven (Corvus
corax) attempted to land near the corpse 2 min later,
but was unable to do so. The next day a raven was
eating the dead bird. On 4 April, another day of strong
north winds with sudden gusts, McNicholl observed
another adult Great Cormorant to be blown about
30 m along the cliff-face on attempting to land. This
bird was able to land briefly at the new locale along the
cliff, but was driven away by other birds there. These
observations were made during regular checks of the
colony from a cleft in the cliff which hid the observer
from birds not actually perched on ledges on the cliff.
This cleft was over 200 m from the spot where the bird
was killed. McNicholl visited the colony from 22
March to 22 April during which period cormorant
activity declined markedly on days with gusty winds,
but birds were able to land readily in high winds if
these were steady.
Byrd and Tobish (1978) attributed the deaths of 48
chick and two adult kittiwakes ( Rissa brevirostris and
R. tridactyla) to wind during a violent storm in
Alaska, but did not observe the precise manner of
deaths. There are many reports in the literature of
deaths of birds during storms, and several reports of
deaths by collison with man-made objects (see review
by Weir 1976), but we are not aware of other reports of
deaths caused by collisions at familiar colony sites in
winds not associated with violent storms. The above
observations suggest that strong gusty winds could
pose a serious problem to large seabirds at cliff
colonies. Stonehouse (1964) mentioned that frigate-
birds (Fregatidae) have difficulty in landing during
strong winds, but did not attribute deaths to this
cause.
These observations were made during a study by
Hogan under the supervision of R.D. Morris.
Financial support was provided by the Prince Edward
Island Fish and Wildlife Division of the Department
of the Environment, a National Research Council of
Canada grant to R.D. Morris, and an Ontario
Graduate Scholarship to Hogan. The manuscript was
improved on the basis of comments by R. K. Ross and
an anonymous reviewer.
Literature Cited
Byrd, G. V. and T. G. Tobish. 1978. Wind-caused mortali-
ty in a kittiwake colony at Buldir Island, Alaska.:
Murrelet 59: 37.
Stonehouse, B. 1964. Frigatebird. Jn A new dictionary of
birds. Edited by A. L. Thomson. McGraw-Hill, New
York. pp. 325-326.
Weir, R. D. 1976. Annotated bibliography of bird kills at
man-made obstacles: a review of the state of the art and
solutions. Canadian Wildlife Service, Ontario Region,
Ottawa.
Received 18 September 1978
Accepted 17 November 1978
176
THE CANADIAN FIELD-NATURALIST
Vol. 93
Habitat Selection by Wintering Snowy Owls (Nyctea scandiaca)
M. Ross LEIN and G. A. WEBBER
Department of Biology, University of Calgary, Calgary, Alberta T2N 1N4
Lein, M. Ross and G. A. Webber. 1979. Habitat selection by wintering Snowy Owls (Nyctea scandiaca). Canadian Field-
Naturalist 93: 176-178.
Habitat selection by Snowy Owls (Nyctea scandiaca) wintering near Calgary, Alberta, was studied in 1973-1976. Habitats
were classified as being residential, fallow, stubble, hayfield, or pasture, and areas of each were determined. Compared to the
relative areas of the various habitats, owls were seen more frequently in stubble and hayfields, and less frequently in the other
three habitat categories. This selection is believed to be related to the varying densities and accessibility of small mammals in
the different habitats.
Key Words: Snowy Owls, habitat selection, winter, prairie farmland, prey availability.
Studies of the behavioral ecology of the Snowy Owl
(Nyctea scandiaca) in winter are sporadic and
incomplete. Gross (1927, 1931, 1944, 1947) docu-
mented several irruptions between 1926 and 1946
but did not obtain comprehensive information on
aspects of the owls’ ecology other than food habits.
Most subsequent publications deal with unusual
locality records or numbers of birds, or with food
habits, although a few (Keith 1960, 1964: Quilliam
1965: Weir 1973) consider other behavioral or
ecological aspects. The virtual lack of intensive or
long-term investigations of the biology of the species
in areas where it regularly winters is especially
Surprising since some owls may actually spend more
time in the south than on their arctic breeding grounds
(Bird 1972).
The senior author began a long-term study of
wintering Snowy Owls near Calgary, Alberta in 1973.
Snowy Owls have been recorded in numbers in this
area every winter since 1964-65 (Bird 1972: personal
observations). This note reports some initial findings
of this investigation on the habitats utilized by owls on
their winter range.
Methods
Each winter from 1973-74 to 1975-76, 8-12 survey
routes were censused by members of the Calgary Field
Naturalists’ Society, using a modification of Gollop’s
(1965) procedure. Each 64-km (40-mi) route was
surveyed every 2-4 wk by two volunteers who drove
the route at a speed of 30-50 km/h, stopping every
3.2 km (2.0 mi) to scan the surrounding area with
binoculars. Each owl’s location was plotted on a
topographic map and pertinent standardized data,
including a habitat description, were recorded.
This analysis is limited to one route in 1974—75 and
three routes in 1975-76 because of the effort involved
in calculating the habitat composition of routes and
the availability of recent aerial photography. The
routes chosen were the most productive in terms of
total owls recorded, and were located in mixed
agricultural land, just east of Calgary.
A habitat map was prepared for each route,
including an area extending 1.6 km (1.0 mi) on each
side of the survey route and 1.6 km before the
beginning and after the ending of the route. The
nature of each field or other habitat unit was identified
from aerial photos or by direct inspection. All habitat
determinations were made after cultivation ceased in
the fall and prior to its resumption in the spring. Areas
of each habitat on each route were estimated using the
standard dot grid method.
We recognize five habitat types on the study area:
1. Residential: farmyards, rural residences, feedlots,
hamlets, and other areas of high human activity.
2. Summerfallow: cultivated fields where either no
crop was planted during the previous growing season
or the stubble was ploughed up after harvest. These
areas are virtually bare earth with only traces of plant
material on the surface.
3. Stubblefields: areas where cereal crops have been
grown and harvested, leaving only the basal 10-20 cm
of the stems. Fields usually persist in this form until
ploughing and seeding in the following spring.
4. Hayfields: areas planted to non-native grasses and
cut yearly for livestock fodder. They are rarely used as
pasture and are ploughed and reseeded at intervals of
longer than one year.
5. Pastures and Sloughs: areas not subjected to
annual cultivation for the raising of cereal crops or
hay, often marginal land which may be used as pasture
for livestock. Sloughs fill with spring runoff but are
typically dry throughout much of the summer,
autumn, and winter. This is a highly heterogeneous
category which is not easily subdivided.
We tallied the survey results by the habitat in which
-each owl was located when sighted. When an owl was
perched on a fenceline that represented a boundary
IA)
between two different habitats, a value of 0.5 owl was
assigned to each of the two categories.
Results
The distribution of habitat classes on the survey
routes is presented in Table 1. The major category is
stubblefields, comprising almost 60% of the area.
Residential areas are a very minor category.
The distribution of owl sightings across these
habitat classes is also shown in Table |, along with an
expected distribution of sightings based on a null
hypothesis of no habitat selection. The observed
distribution is significantly different from that expec-
ted (chi-square goodness of fit test, P< 0.01), demon-
strating a degree of habitat selection. Owls are
recorded more frequently than expected in stubble
and hayfields, and less frequently than expected in the
other three habitat categories.
Discussion
The overall preference for open habitats by winter-
ing Snowy Owls is well-known (Bent 1938: Gross
1947: Keith 1960). In the Calgary area, most owls are
found in agricultural land east of the city: few birds are
observed in the more wooded aspen parklands or
foothills to the west (Bird 1972; M.R. Lein, un-
published data). A similar avoidance of wooded
regions is noted elsewhere (Gross 1947: Nagell and
Frycklund 1965); however, the present study appears
to be the first attempt to demonstrate habitat selection
by Snowy Owls at a finer level.
We suggest that the preference for stubblefields and
hayfields is related to the availability of small-
mammal prey in such areas, as would be predicted by
optimal foraging theory (MacArthur and Pianka
1966; Pyke et al. 1977). Summerfallow fields are
relatively barren and offer little to rodents either in
cover or food. Pastures are usually heavily grazed and
offer little cover for small mammals. Non-grazed
sloughs and roadside ditches offer excellent cover for
rodents, which are relatively abundant in such
habitats (P. C. Boxall, unpublished data), but rank
vegetation and deep snow cover may sharply limit
NOTES 177
their availability to owls. Low numbers of owls
around residential areas may be a direct response to
human disturbance. Thus, the habitat distribution of
the owls suggests that they may be choosing areas
(stubblefields and hayfields) in which rodents are both
relatively numerous (P. C. Boxall, unpublished data)
and available to predators.
It is perhaps surprising that a more pronounced
degree of habitat selection is not shown. Several
reasons may be suggested. The first is the relatively
crude level of the analysis. Owls certainly do not
categorize habitats in the same manner as do human
investigators, and an analysis of distribution using
different habitat classes might show more striking
differences. Second, and perhaps more important, is
the implicit assumption of the present study that the
owls recorded in a habitat were actually choosing it as
a hunting area. Although hunting attempts have been
observed in all habitats throughout the day, it is
possible that owls may choose different areas for
roosting and “loafing” than those in which they hunt.
We know very little about the hunting behavior or
activity cycles of wintering Snowy Owls. Owls tend to
use elevated perches such as trees or utility poles early
in the morning and toward sunset (Keith 1964: M. R.
Lein, unpublished data) and to perch on or near the
ground during mid-day. It has been suggested that
these high perches are hunting perches and that this
represents a diurnal pattern of hunting behavior.
Unfortunately, most of the data used in this study
were gathered during the middle of the day and are
unsuitable for a test of such an hypothesis. If it were
possible to identify owls likely to be actively hunting,
and to carry out the analysis on only those owls,
stronger habitat selection might be expected.
Acknowledgments
We thank those members of the Calgary Field
Naturalists’ Society who conducted owl surveys: this
study would not have been possible without their
efforts. We gratefully acknowledge P. C. Boxall for
providing us with preliminary results of his investiga-
tion on Snowy Owls, and for commenting on the
TABLE |—Distribution of habitats, and of owl sightings, on the survey routes analyzed.
Habitat category
Pasture
Residential Fallow Stubble Hayfield + slough Total
Percentage of total area 1.81 Da 58.48 6.55 18.04 100.00
Number of owls observed 2.5 20.5 116.0 18.0 18.0 175.0
Number of owls expected Ball) 26.46 102.34 11.46 SiESy/ 175.00
178 THE CANADIAN FIELD-NATURALIST
manuscript. T. E. Dickinson also provided helpful
suggestions. Financial support was received in the
form of operating grants from the University of
Calgary and the National Research Council of
Canada to M. R. Lein.
Literature Cited
Bent, A. C. 1938. Life histories of North American birds of
prey. Part 2. United States National Museum Bulletin
Number 170.
Bird, C. D. 1972. The Snowy Owl in the Calgary area,
1964-1972. Calgary Field Naturalist 3: 106-107.
Gollop, J. B. 1965. Snowy Owls and Gray Partridge near
Saskatoon in the winter of 1963-1964. Blue Jay 23:
28-31.
Gross, A. O. 1927. The Snowy Owl migration of 1926-27.
Auk 44: 479-493.
Gross, A. O. 1931. Snowy Owl migration — 1930-31. Auk
48: SOI-S11.
Gross, A. O. 1944. Food of the Snowy Owl. Auk 61: I-18.
Gross, A. O. 1947. Cyclic invasions of the Snowy Owl and
the migration of 1945-46. Auk 64: 584-601.
Vol. 93
Keith, L. B. 1960. Observations on Snowy Owls at Delta,
Manitoba. Canadian Field-Naturalist 74: 106-112.
Keith, L. B. 1964. Territoriality among wintering Snowy
Owls. Canadian Field-Naturalist 78: 17-24.
MacArthur, R. H. and E. R. Pianka. 1966. On optimal use
of a patchy environment. American Naturalist 100: 603-
609.
Nagell, B. and I. Frycklund. 1965. Invasionen av fjalluggla
(Nyctea scandiaca) i sodra Skandinavien vintrarna 1960-
1963 samt nagot om artens beteende pa Overwintring-
slokalerna. Var Fagelvarld 24: 26-55.
Pyke, G.H., H.R. Pulliam, and E. L. Charnoy. 1977.
Optimal foraging: a selective review of theory and tests.
Quarterly Review of Biology 52: 137-154.
Quilliam, H.R. 1965. Winter study of Snowy Owls on
Wolfe Island, 1965. Ontario Field Biologist 19: 1-8.
Weir, R. D. 1973. Snowy Owl invasion on Wolfe Island,
winter 1971-72. Ontario Field Biologist 27: 3-17.
Received 21 March 1978
Accepted 17 November 1978
Two-lined Salamander, Eurycea bislineata, in Labrador
FRANCIS R. COOK! AND JOHN PRESTON2
'Herpetology Section, National Museum of Natural Sciences, Ottawa, Ontario KIA 0M8
2Environment Canada, 814 Hudson Drive, Labrador City, Newfoundland A20O I1M5
Cook, Francis R. and John Preston. 1979. Two-lined Salamander, Eurvcea bislineata, in Labrador. Canadian Field-
Naturalist 93(2): 178-179.
In mid-August 1972 a single Two-lined Sala-
mander, Eurycea bislineata, was collected by J. P.
approximately 16 km (10 mi) NE of Labrador City on
the west side of a small brook that drains into Moose
Head Lake, about 0.6 km (1 mi) upstream from the
lake (52°58’N, 66°40’W). The specimen is a recently
transformed juvenile 63mm in total length and
29.5mm snout to anterior margin of vent. It is
catalogued as National Museum of Natural Sciences
(Herpetology Section) 17948.
This is the first example of the species collected in
Labrador and is a 290-km (180-mi) range extension
north of the nearest records on the north shore of the
Gulf of St. Lawrence (Bleakney 1958, map 85: Logier
and Toner 1961, map 19; Power 1965). Based on two
collections which have not been previously docu-
mented, Conant (1975, map 240) shows that in
interior central Quebec the range of the Two-lined
Salamander extends north to near the latitude of
southern Labrador. These collections are a single
specimen from the Lac Charon area 32 km (20 mi)
from Chibougamau taken by Jean-Guy Rossignol 3
July 1967 (NMNS 10287), and five specimens col-
lected along the Rupert River, 48 km (30 mi) west of
its source at Lake Mistassini approximately 20 July
1967 by Daniel Roy (American Museum of Natural
History 88025-29; transmitted by Roger Conant).
Bleakney (1968, map 82) plotted a record of the Red-
backed Salamander, Plethodon cinereus, in the Lake
Melville area of central Labrador. This was based ona
report from W. J. Smith who in the summer of 1955
had the local occurrence of a small salamander
mentioned to him on several occasions (Bleakney
1958, p. 15). Because of gross similarities between
Plethodon and Eurycea, however, we suggest that this
record may be more logically assigned to the latter and
1979
suggest a wider Labrador range for it than our single
specimen establishes. The presence of Eurycea in
Labrador and central Quebec is a further indication of
the need for a northward revision of Bleakney’s
Herpetofaunal Section 5 to include his previously
disjunct Section 5A as has already been proposed by
Cook and Folinsbee (1975) on the basis of recent
records for the Blue-spotted Salamander, Amby-
stoma laterale, the only other salamander verified for
Labrador to date.
Literature Cited
Bleakney, J. Sherman. 1958. A zoogeographical study of
the amphibians and reptiles of eastern Canada. National
Museum of Canada Bulletin 155: 1-119.
NOTES 179
Conant, Roger. 1975. A field guide to reptiles and am-
phibians of eastern and central North America. Houghton
Mifflin Company, Boston. xviii + 429 pp.
Cook, Francis R. and John Folinsbee. 1975. Second record
of the Blue-spotted Salamander from Labrador. Canadian
Field-Naturalist 89(3): 314-315.
Logier, E. B.S. and G. C. Toner. 1961. Check list of the
amphibians and reptiles of Canada and Alaska. Royal
Ontario Museum, Toronto, Life Sciences Division Con-
tribution 53: 1-92.
Power, G. 1965. Notes on the cold-blooded vertebrates of
the Nabisipi River region, County Duplessis, Quebec.
Canadian Field-Naturalist 79(1): 49-61.
Received 9 November 1978
Accepted 4 December 1978
First Canadian Record of a Flathead Catfish
E. J. CROSSMAN! and J. H. LEACH?
‘Department of Ichthyology and Herpetology, Royal Ontario Museum, Toronto, Ontario M5S 2C6
2Ontario Ministry of Natural Resources, Fisheries Research Station, RR #2, Wheatley, Ontario NOP 2P0
Crossman, E. J. and J. H. Leach.
179-180.
1979. First Canadian record of a Flathead Catfish. Canadian Field-Naturalist 93(2):
Pylodictis olivaris was captured in the Canadian waters of Lake Erie on 22 August 1978. This is a 30-km range extension north
and is the first Canadian record.
Key Words: Flathead Catfish, Pv/odictis olivaris, Canadian record, Lake Erie.
A live specimen of Pylodictis olivaris, the Flathead
Catfish (Ictaluridae), was netted in Lake Erie and
constitutes the first time the species has been captured
in Canadian waters.
The fish was taken by Rudy Krause, of K. W.
Krause and Sons Fisheries, Leamington, Ontario, on
22 August 1978. It was caught ina commercial trapnet
west of Point Pelee, 4.8 km north of the tip of the
point (41°56’50”N, 82”31’40”W). The bottom of the
net site was silty clay and the water temperature at the
time of capture was 24.3°C.
The catfish was 52 cm in total length and weighed
1.79 kg. Trautman (1957) indicated that the species
has been known to reach 134.6 cm length and 37.2 kg
weight. The Canadian specimen had a noticeable
deformity of the middle region of the caudal verte-
brae, but otherwise agreed with the morphological
characteristics of the species as given by Trautman
(1957) and Hubbs and Lagler (1964). The most
striking feature other than head shape, which separ-
ates this species from other local catfishes, is the
strong backward extensions of the premaxillary
tooth-patches. These were very obvious in this
specimen.
The species has long been known in the United
States waters of Lake Erie, and Trautman (1957)
indicated records north to the international boun-
dary. This latest record from the northern side of Lake
Erie itself represents a range extension of 30 km north
of previous records and is somewhat unusual because
this species is more abundant in the Ohio River and its
tributaries than in the Lake Erie portion of Ohio.
Trautman listed only five specimens taken in the Lake
Erie waters of Ohio from 1938 to the date of writing
(1956?). He suggested that “there appears to be a small
population . . . in the Huron River, where they are
taken yearly.” The mouth of the Huron River in Ohio
is due south of Point Pelee.
In describing the total range of the species, Hubbs
and Lagler (1964) called it rare in Lake Erie and
mentioned that it is usually a large-river form. This
catfish had not been previously reported from
180
adjacent Canadian waters, in spite of the intensive
commercial fishery in that area. It was the only
specimen seen so that it is impossible to say now
whether it is a stray or whether a small population has
already been established in the Point Pelee area.
The shallow waters from western Lake Erie to Lake
St. Clair have contributed most of the limited number
of records of freshwater fishes that have moved north
into Canadian waters over the past 25 yr. Catches in
those waters should be carefully scrutinized and all
unusual items reported.
The specimen has been added to the reference
collection of the Royal Ontario Museum (ROM Cat.
No. 34561).
THE CANADIAN FIELD-NATURALIST
Vol. 93
We thank Rudy Krause for his care in recognizing
that the fish was different, and for taking the trouble
to turn it over to the Ontario Ministry of Natural
Resources.
Literature Cited
Hubbs, C. L. and K. F. Lagler. 1964. Fishes of the Great
Lakes Region. University of Michigan Press, Ann Arbor,
Michigan. 213 pp.
Trautman, M.B. 1957. The fishes of Ohio. Ohio State
University Press, Columbus, Ohio. 683 pp.
Received 2 October 1978
Accepted 4 December 1978
Earthworm Cocoons as a Drift Component in a
Southern Ontario Stream
DONALD P. SCHWERT! and KENNETH W. DANCE?
'Geology Department, Stevens Hall, North Dakota State University, Fargo, North Dakota 58105
2Ecologistics Limited, 309 Lancaster Street West, Kitchener, Ontario N2H 4V4
Schwert, D. P. and K. W. Dance. 1979. Earthworm cocoons as a drift component in a southern Ontario stream. Canadian
Field-Naturalist 93(2): 180-183.
Cocoons representing at least six species of Lumbricidae were isolated from drift subsamples of a stream in Waterloo County,
Ontario in the wpring of 1976. Of the 308 cocoons, 92% were viable. We present arguments to support cocoon drift as a
potentially important mechanism in lumbricid dispersal.
Key Words: cocoons, Lumbricidae, Oligochaeta, biological drift, dispersion, Ontario, geographical distribution.
In southern Ontario, as in many other regions of
North America, large populations of lumbricid
earthworms often occur in moist lowlands adjacent to
streams and lakes. When temperature, moisture, and
light conditions permit, a number of the lumbricid
species in these areas are active on or just below the
soil surface. Significant activity combined with
surface runoff from rainstorms and snowmelt can
result in large numbers of earthworms and their
cocoons becoming trapped in waterways. Their
subsequent downstream drift might be expected to be
a significant factor in lumbricid dispersal, but this
mechanism has remained poorly studied. Bouché
(1972) hypothesized that stream drift had an impor-
tant influence on the distribution of some hygrophilic
lumbricid species in France, and Ward (1976) applied
a similar hypothesis to explain the recolonization of a
riffle area by the lumbricid Evseniella tetraedra in a
Colorado stream. No published observations of the
drift of lumbricid cocoons exist, however, probably
because these might be easily confused with plant
seeds during sorting.
During the latter 5 mo of a 13-mo stream drift study
in southern Ontario (Dance and Hynes, in press),
sorting of earthworm cocoons from freshly collected
drift samples was initiated. We present here data on
this component and suggest its possible significance
with respect to lumbricid dispersal.
The Study Area
All of the examined cocoons were obtained from
drift samples of two headwater branches of Cana-
gagigue Creek in a predominantly agricultural region
immediately north of Floradale in Waterloo County,
Ontario (43°39’N, 80°35’W). Stations | and 2 were
located on the permanent, spring-fed east branch of
ITY
the creek. Stations 3 and 4 were located on the west
branch, an intermittent stream which flowed continu-
ously during the five months of this study. Station |
was a gravelly riffle, in a treeless pasture 350 m
downstream from a White Cedar ( Thuja occidentalis)
woods. Station 2 was a cobble and boulder riffle at the
downstream margin of an extensive American Beech
(Fagus grandifolia) and Sugar Maple (Acer sac-
charum) stand. Station 3 was at the downstream edge
of a short, silty riffle in a treeless pasture. The
streambank at Station 4 was lined with herbs and
Salix spp., and the bed consisted of boulders and
cobbles resting on silts.
Methods
Drift was subsampled continuously using a special-
ly designed apparatus of 253 um Nitrex cloth net
mounted over an aluminum and steel frame (Dance et
al., in press). Water flowed without wave interference
through a vertical opening | cm wide and 100 cm high
into a long sample bag. The contents of the trap were
placed weekly into a plastic bag and frozen until
sorting. Under a binocular microscope, cocoons were
sorted from thawed samples which had been collected
between February and June 1976 and placed into
labelled vials of 70% ethanol.
Attempts at placing generic or specific names on the
cocoons were inhibited by distortions in the cocoon
shape resulting from the sample procedures and by the
frequent absence of the diagnostic end “tufts” which
had apparently broken off during the turbulence of
drift. Names could, therefore, be placed on only a
small number of the cocoons through comparison
with a cocoon reference collection and with the
cocoon descriptions in Evans and Guild (1947) and
Gerard (1964).
Mean monthly discharge volumes during the study
period for each of the sample stations were calculated
from data on the creek provided by the Canada Water
NOTES
18]
Survey and the School of Engineering at the Uni-
versity of Guelph. Utilizing this information and the
calculated average component of the total discharge
continuously subsampled by each trap (for detailed
procedure, see Dance et al., in press), a multiplication
factor was derived estimating the total monthly
number of cocoons passing each sample point.
Results
A summary of the mean monthly discharge values
for the four Canagagigue Creek sample stations is
presented in Table 1. Discharge volumes fluctuated
widely for each station during the study period but for
all four were maximal in March and minimal in June.
Monthly summaries of the actual cocoon content of
the drift samples and of the estimated total cocoons
flowing past each station are presented in Table 2.
Cocoons were absent from all February samples, but
maximum numbers were recorded in March at the
east branch stations and in April at the west branch
stations; the west branch stations were inoperable
during most of March because of flooding. Except for
a sharp increase in June numbers at Station 2, the
incidence of cocoons generally decreased into the
early summer.
TABLE |—Estimated mean 1976 monthly discharge (L/s)
for four Canagagigue Creek drift sample sites (Waterloo
County, Ontario)
Station
Month | 2 3 4
Feb. 202 455 3174 3398
Mar. 901 1126 3066 6089
Apr. AY || 410 394 446
May 151 275 220 298
June 148 24] 47 87
TABLE 2—Summary of the 1976 monthly totals of earthworm cocoons per drift trap (and estimated total cocoons flowing
past each sample station) for four Canagagigue Creek drift sample sites in Waterloo County, Ontario, and their percent
viability
Number of cocoons (estimated total)
Station Numbers of viable
Cocoon cocoons (% of
Month | 2 3 4 total monthly total)
Feb. 0 (0) 0 (0) 0 (0) 0 (0) 0 = (=)
Mar. 52 (17731) 42(22 105) Flood Flood 94 88 ( 93.6)
Apr. 34 ( 5964) 33 ( 9167) 10 (1149) 52 (11 556) 129 110 ( 85.3)
May 22 ( 4000) 6 ( 1875) 1( 88) 8 ( 2963) 37 36 ( 97.3)
June 18 ( 3529) 29 (13 809) 0( 0) 1( 345) 48 48 (100.0)
Total 308 282 ( 91.8)
182
Nearly 92% of the cocoons were found to contain
sperm and albumen, or some stage in the develop-
ment of the embryonic mass, and these were termed
“viable,” or potentially capable of hatching.
All cocoons resembled in general form those of the
Lumbricidae. A qualitative examination of the undis-
torted component yielded cocoons resembling the
following taxa in the family: Aporrectodea spp.,
Dendrodrilus rubidus, Eisenia fetida, Eiseniella tet-
raedra, Lumbricus terrestris, Octolasion teyrtaeum.
We have collected adults of each of the above
species from sites in Waterloo County, and all of the
species have been recorded from southern Ontario by
Reynolds (1977). One adult Eisenia fetida and one
adult Eiseniella tetraedra were also collected from the
drift samples.
Discussion
Fluctuations in the monthly totals of the observed
cocoons undoubtedly reflect both the physical state of
the environment and the cocoon production of the
local fauna. The absence of cocoons from the
February samples would be expected because of the
frozen condition of the soil, but the March and April
cocoon peaks are probably the result of both the high
rate of surface runoff from snowmelt and rain and a
high degree of surface activity and cocoon pro-
duction by the earthworm fauna. The precise dis-
tances which any of the cocoons drifted to the trap
sites remains unknown, although presumably none of
the cocoons were derived from sources greater than
2-3 km upstream, the area of the upper Canagagigue
watershed.
The small size (< 6mm) and tough, spheroidal
outer walls of lumbricid cocoons are ideally suited for
long and rigorous transport in streams. Individuals of
several lumbricid species have been shown to be
capable of prolonged submersion (Roots 1956;
Edwards and Lofty 1972), and the successful hatching
and growth while submerged of the lumbricid
Allolobophora chlorotica has been demonstrated by
Roots (1956). From these studies and from the
remarkably high viability of the drift cocoons of the
present study, successful hatching of cocoons could be
expected in areas of a stream where the cocoons had
been deposited near the margin or in the bottom
sediments of pools. A major consequence, therefore,
would be the establishment of the transported species
in lowland areas along the stream.
Because cocoons have subsequently been obtained
from random drift samples in other streams of
southern Ontario and of Newfoundland, the phe-
nomenon of cocoon drift appears to be widespread.
We have also collected large immature and adult
Lumbricidae from other streams in Ontario, as wellas
in Newfoundland, New York, and Pennsylvania.
THE CANADIAN FIELD-NATURALIST
Vole9s
Although these worms survived submergence in
rigorous environments, the battered nature of many
of the specimens indicates that these large, soft-bodied
stages have difficulty in physically surviving pro-
longed transport. In addition, this stage is particularly
susceptible to predation by fish; the palatability of
cocoons to fish is unknown, but presumably low.
The successful drift of cocoons may be of con-
siderable importance in explaining the modern dis-
tribution of many lumbricid species in North Ameri-
ca. All of the species identified from Canagagigue
Creek are of taxa hypothesized by Gates (1970) and
Reynolds (1974) as having been introduced into North
America from Europe. The widespread distribution of
many of these species across Canada and the United
States is attributed by Gates (1976) as being primarily
due to transport by human activity. While man has
undoubtedly influenced earthworm distribution, he
cannot, as Ball (1975) argues, logically claim respon-
sibility for the entire distribution of lumbricid
earthworms across the continent. Transport of co-
coons, however, from one point of intensive lumbricid
establishment upstream to other points downstream
could result in the subsequent colonization of the
lower watershed by these species. Although further
investigation is needed to support this mechanism,
stream drift may be as important to the dispersal of
earthworms as it is to many other groups of
invertebrates.
Acknowledgments
We are grateful to those property owners near
Floradale, Ontario, who granted us access to Cana-
gagigue Creek for sampling. We also appreciate the
useful suggestions of A. D. Harrison of the University
of Waterloo regarding the manuscript. Funding for
publication of this paper was provided by Ecologistics
Limited and by North Dakota State University.
Literature Cited
Ball, I. R. 1975. Nature and formulation of biogeographi-
cal hypotheses. Systematic Zoology 24(4): 407-430.
Bouché, M. B. 1972. Lombriciens de France, écologie et
systématique. Institut National de la Recherche Agrono-
mique, Paris. 617 pp.
Dance, K. W. and H. B. N. Hynes. /n press. A continuous
study of the drift in adjacent intermittent and permanent
streams. Archiv fur Hydrobiologie.
Dance, K. W., H. B.N. Hynes, and N. K. Kaushik. /n
press. Seasonal drift of solid organic matter in two
adjacent streams. Archiv fur Hydrobiologie.
Edwards, C. A. and J. R. Lofty. 1972. Biology of earth-
worms. Chapman and Hall, Ltd., London. 283 pp.
Evans, A.C. and W.J. McL. Guild. 1947. Cocoons of
some British Lumbricidae. Annals and Magazine of
Natural History, Series I1, 14: 714-719.
Gates, G. E. 1970. Miscellanea megadrilogica VII. Mega-
drilogica 1(2): 1-14.
1979
Gates, G. E. 1976. More on earthworm distribution in
North America. Proceedings of the Biological Society of
Washington 89(40): 467-476.
Gerard, B. M. 1964. Synopses of the British fauna. Number
6. Lumbricidae (Annelida) with keys and descriptions.
Linnaean Society of London. 58 pp.
Reynolds, J. W. 1974. Are oligochaetes really hermaphro-
ditic organisms? Biologist 56(2): 90-99.
Reynolds, J. W. 1977. The earthworms (Lumbricidae and
Sparganophilidae) of Ontario. Life Sciences Miscel-
laneous Publications, Royal Ontario Museum. x + 141
PP.
NOTES
183
Roots, B. I. 1956. The water relations of earthworms. II.
Resistance to desiccation and immersion, and behaviour
when submerged and when allowed a choice of environ-
ment. Journal of Experimental Biology 33: 29-44.
Ward, J. V. 1976. Lumbricid earthworm populations in a
Colorado mountain stream. Southwestern Naturalist
AME Wns.
Received 29 September 1978
Accepted 27 December 1978
Flowering Plant Phenology at Sheep Mountain, Southwest
Yukon Territory
MANEFRED HOEFS
Yukon Game Branch, P.O. Box 2703, Whitehorse, Yukon Territory YIA 2C6
Hoefs, M.
93(2): 183-187.
1979. Flowering plant phenology at Sheep Mountain, southwest Yukon Territory. Canadian Field-Naturalist
The dates of initiation of flowering of 60 vascular plant species in the Kluane Lake area, Yukon Territory, are presented for the
1970, 1971, and 1972 seasons. The altitudinal advance of plant development is determined, using Carex filifolia as the
indicator species.
Key Words: phenology, flowering plants, subarctic botany, Carex filifolia, Kluane Lake, Yukon Territory.
The data presented here are part of a larger
investigation carried out on Dall Sheep (Ovis dalli
dalli) and their range in the Kluane Lake area of the
Yukon Territory since 1969. The specific study area is
“Sheep Mountain,” an important winter range of a
Dall Sheep population located at the southeast shore
of Kluane Lake near the mouth of the Slims River
(61°00’N, 138°30’E). For details on the vegetation,
climate, geology and soil of the area, the reader 1s
referred to Hoefs et al. (1975) and for details on the
sheep population, their range use patterns, and forage
selection, to Hoefs (1975).
During this investigation it became clear that a
number of activities of sheep, for instance forage
selection, use of various plant communities, and
vertical migration, were closely linked to plant
phenological phenomena. A number of these were
investigated.
My studies (Hoefs 1974) have demonstrated that
the flowers of certain plant species are preferred
forage items for Dall Sheep. This paper deals with the
dates of initiation of flowering of 60 vascular plants on
Sheep Mountain.
A number of factors have been used to explain the
vertical seasonal movements of ungulates; these
include snow conditions and other weather factors,
avoidance of blood-sucking insects, protection of
winter ranges, and advantages with respect to forage
(Dixon 1938; Murie 1944: Blood 1963; Egorov 1967;
Hebert 1972). Some indications of the vertical march
of phenology can be obtained from the dates of
first flowering of plant species at different altitudes. A
more accurate quantification, however, is possible by -
observation of the performance of the same species of
plant at various altitudes. This study determined
altitudinal advance using Carex filifolia as the
indicator species. Carex filifolia was selected because
it is (a) a fairly abundant plant in all dry grassland
associations in the boreal, subalpine, and alpine
biogeoclimatic zones of the study area (Hoefs et al.
1975); (b) one of the most preferred Dall Sheep forage
plants (Hoefs 1975); (c) an “early bird” in a pheno-
logical sense, flowering earlier and reaching the
annual maximum of growth before any of the other
important forage plants (Figure 1).
Methods
Sixty species of flowering plants, known to be used
by sheep (Hoefs 1975), were mapped and marked by
wooden stakes on Sheep Mountain in 1969. The dates
184
THE CANADIAN FIELD-NATURALIST
Species April May June July
20 30 10 20 30 10 20 30 10 20
Pulsatilla patens mm | | ee
Carex filifolta ee meee | | ee
Townsendta hookertr | fees Rae |
Arctostaphylos rubra i ie a ae eee Ee
Anemone parviflora eee | eae ee ee ee
Potentilla hookertana eng | ee
Saxtfraga opposttifolta Rat (eee eae)
Oxytropts visetda oe | Tees | S| | ea
Lupinus arcticus | me ee
Arctostaphylos uva-urst a aaa
Linum perenne = Ce ed
Erigeron composttus ae SS Lr aa)
Plantago canescens aS an ae ea ee ee a
Smilacina racemosa Sees | | pat le)
Carex aquatilis mT Se een re
Hedyearum alpen). Tas Fee | ee ae |
Amevanchversaint porta) (ee Se ee |
Mertensta paniculata a er eee ee
OCNSEC TECTED ESS ae. |. 2 le
wnemoneimelte peda BN 2 |
Anemone drummondtr ee] ee eee eee eee
Dryas drummondit [| | fT
Saxtfraga tricusptdata aaa ae
Dryas integrifolia [| oot TT
Casstope tetragona ea ae eS
Taraxacum officinale =r SS
pilopium latifoltum ———==rS
Astragalus umbellatus a aE ee See
Lappula myosotts an ee ee eee ee
Potentilla fruttcosa Pe ee
Myosotts alpestris eee cee
Aster stbtrtcus |. eel
Pentstemon gormant an ae ee ee ee eee
Erigeron purpuratus [ee ee ee
Aster alptnus i a
Oxytropts huddeTsonit i as ee ee
Sedum rosea PI ao
Rosa actcularts LO a ee ee
tlene acaults esac Sipe ae. a as
Artemtsta hyperborea ae Se haar area
Arntea alptna
Vieburnum edule
Aehtllea borealts
Artemista alaskana
Itnnaea borealts PeSaeS h O| nz oe Cie)
pi lobiummuanguetd folie 0 | ee
Calamagrostts purpurascens Poe eee
Vaeetntum vttts-tdaea
Aster yukonense
Ledum palustre
Poa glauca
gadenus elegans
oltdago multrtradiata
Crepte nana | || dl lo
DEC CNIS UCI feta GC CLC a | oe Ol | i |
Erigeron caespitosus ecese| Ge ss | |e 5| ele |
altum boreale (ce a ae a a ay ESTs |
Hordeum jubatum Pe ee ie ae ein a a
Artemisia rupestris aie a eS ee Lz 2
Agropyron yukonense
FIGURE |.
Dates of first flowering of Sheep Mountain plants for 1970, 1971, and 1972.
Vol. 93
179
at which these plants first developed fully opened
flowers were recorded during 1970, 1971, and 1972
seasons.
Ten clumps of Carex filifolia were selected at each
of five altitudinal sites (935, 1000, 1330, 1590, and
1910 m) and measurements of growth, flowering,
fruiting, and initiation of dormancy were carried out
at weekly intervals from early May to late October
1970. The selected clumps were protected from
grazing by exclosure fences.
Nomenclature of plant species follows Hultén
(1968) with the exception of Artemisia hyperborea,
which is described in Porsild (1966).
Results and Discussion
Dates of First Flowering
Figure | lists 60 flowering plants that grow on the
range of the Dall Sheep population under study, and
gives their first flowering dates arranged approxi-
mately in chronological sequence. Because observa-
tions were carried out over three seasons, with
considerable differences in weather patterns, the first
flowering dates are ranges of 10d to 2 wk and not
individual days. Great variations were observed
particularly between the spring of 1971 compared to
1970. Some flowers came into bloom as much as 12 d
earlier in 1971. The data of Figure | can therefore be
interpreted as follows: beginning of range is initial
flowering date in 1971, end of range is initial flowering
date for 1970. The date for 1972 is about half-way
between those two extremes. The data presented here,
particularly for June when many plants came into
bloom, must be considered as approximations since it
was not physically possible to check every marked
species every day. It is unlikely, however, that any of
the dates are in error by more than 4d.
Figure | includes a number of species which grow in
various altitudes and in various plant associations
(aspects) at the same altitude. Even though only
specimens growing in the most “advanced” sites in a
phenological sense, which were usually south-facing
slopes in the boreal zone, were marked and are
considered in Figure I, records were kept on the
delays observed if such species also grew on north-
facing slopes and at higher elevations. Detailed
information on the floristic composition of the
various plant associations on Sheep Mountain, and
their distribution with respect to aspect and altitude
are given in Hoefs et al. (1975) and need not be
repeated here. In general it was found that there was a
10- to 14-d delay in plant development between north-
and south-facing slopes at the same altitude, and a
delay of about | d for each 35 m of altitude at the same
aspect. An example may demonstrate this point.
Pulsatilla patens is the first plant, with the exception
of willows, to have open flowers. On sunny, south-
NOTES 185
facing protected slopes with grassland vegetation in
the boreal zone (1000 m), its flowers may in favorable
springs (1971) be seen as early as 22 April. This species
grows also in alpine elevation (1667 m), where it did
not come into bloom before 17 May (1971). Figure |
reveals that Pulsatilla patens is the only species that
may come into bloom in April, about 26 species begin
to bloom during May in an average year, 19 species in
June, and 15 species in July. Although there are many
exceptions to this rule it is in general true that
members of the families Ranunculaceae and Legu-
minosae are among the first to come into bloom while
members of the Compositae and Gramineae families
are among the last.
Altitudinal Advance of Plant Development
Figure 2 shows the growth curves of Carex filifolia
at five sites with increasing altitudes. Lines connecting
the points at which 50% and 100% of the annual
growth was achieved give an indication of the vertical
phenological march. For reasons as yet unexplained,
the plants at an elevation of 1000 m were pheno-
logically more advanced than those at an altitude of
935 m. One reason may have been that the plants at
the lower altitude were more often subject to shade
cast by surrounding trees during the course of a day.
Because the lines connecting 50% and 100% growth
points are parallel it will suffice to discuss one of them.
At an altitude of 1000 m Carex filifolia reached its
maximum annual growth around 18 June, at 1330 m
around 26 June, at 1590 m around 5 July, and near the
peak of Sheep Mountain, at 1910 m, it took till 15
July, 1970.
The speed of vertical advance of plant development
was therefore 910 m per month or about 315 m per
10 d. It is not possible to quantify within similar terms
the sequence of dry-up, since this is influenced not
only by temperature or altitude but also by moisture
deficiency. This speed of vertical advance agrees with
Hopkins’ (1920) bioclimatic law, which states that .
there is a delay in vegetation development of about 3
to 4d for each 100- to 130-m increase in altitude, as
well as with Stoddart and Smith’s (1955) remark that
“Most observers agree that the (altitudinal) difference
approaches one day for each 100 feet of altitude.”
My studies documented a very close relationship
between vertical migrations of Dall Sheep and plant
phenology, particularly in spring. During May and
June sheep leave their winter ranges and move up
high, making use of plant associations with new
growth and avoiding others that are still dormant.
This pattern of range use continues on summer range,
where advanced plant associations on south- and
west-facing slopes were primarily used in June and
early July while plant associations on east and
northern aspects were used in late July and August.
Selection with respect to phenological stage was also
186 THE CANADIAN FIELD-NATURALIST
100
50
100
50
100
50
100
50
% "green'! matter in total biomass
100
50
50 % of annual
_—[—
Vol. 93
growth completed
Maximum of annual growth completed
FIGURE 2. Growth of Carex filifolia at various altitudes in 1970, demonstrating vertical phenological advance.
observed when feeding took place within one plant
association. New growth was preferred to the mature
and dormant stage of a species. Special attention was
paid to flowers, particularly large and showy ones.
Pulsatilla patens and Oxytropis viscida flowers are
important forage items in early spring when the sheep
are still on winter range, while Pedicularis spp., Dryas
integrifolia, Hedysarum alpinum, Epilobium lati-
folium,
and Oxytropis huddelsonii flowers are
important on alpine summer range. Fall migration
from alpine summer ranges commences in late August
and early September. Only ewes and lambs, however,
appear to arrive at lower elevations in time to utilize
certain plant species that are still “green.” Heavy use is
made at this time of the various willow species in the
subalpine shrub zone (1200 to 1600 m), when all the
1979
vegetation of the alpine zone is already dormant.
Forage selection and range use patterns are dealt with
in detail in Hoefs (1975).
Literature Cited
Blood, D. A. 1963. Some aspects of behaviour of a bighorn
herd. Canadian Field-Naturalist 77(2): 77-94.
Egorov, O. V. 1967. Wild ungulates of Yakutia. Israel
program for scientific translations, Jerusalem. United
States Department of Commerce.
Dixon, J. S. 1938. Birdsand mammals of Mount McKinley
National Park. United States Department of Interior,
National Park Service, Fauna Series Number 3.
Hebert, D. 1972. Differences between years and nutrient
cycles. Alberta Fish and Wildlife Division, Northern
Wild Sheep Council, Symposium Proceedings 1972: 15—
DD,
Hoefs, M. 1974. Food selection of Dall sheep. Jn The
behaviour of ungulates and its relation to management.
Volume 2. Edited by V. Geist and F. Walter. Inter-
national Union for Conservation of Nature and Natural
Resources. pp. 759-786.
NOTES
187
Hoefs, M. 1975. Ecological investigation of Dall sheep
and their habitat. Ph.D. thesis, University of British
Columbia, Vancouver. ;
Hoefs, M., I. McT. Cowan, and V.J. Krajina. 1975.
Phytosociological analysis and synthesis of Sheep Moun-
tain, southwest Yukon Territory, Canada. Syesis 8
(Supplement 1): 125-228.
Hopkins, A.D. 1920. The bioclimatic law. Journal of
Washington Academy of Science 10: 34-40.
Hultén, E. 1968. Flora of Alaska and neighboring ter-
ritories. Stanford University Press, Stanford, California.
Murie, A. 1944. The wolves of Mount McKinley. Fauna of
the National Parks of the United States, Fauna Series
Number 5. United States Department of Interior.
Porsild, A. E. 1966. Contributions to the flora of south-
western Yukon Territory. National Museum of Canada,
Contributions to Botany IV, Bulletin 216.
Stoddart, L. A. and A. D. Smith. 1955. Range manage-
ment. McGraw-Hill Book Company, New York. 433 pp.
Received 8 September 1978
Accepted 5 December 1978
Eggshell Thickness in American Shorebirds before and since DDT
MICHAEL L. MORRISON and LLOYD F. KIFF
Western Foundation of Vertebrate Zoology, 1100 Glendon Avenue, Los Angeles, California 90024
Morrison, Michael L. and Lloyd F. Kiff. 1979. Eggshell thickness in American shorebirds before and since DDT. Canadian
Field-Naturalist 93(2): 187-190.
Thickness indices of eggshells collected before and since use of DDT began were obtained for 31 species and subspecies of
American shorebirds. Little change in thickness was found between early and recent samples, the maximum being -4.7% in
“American” Oystercatcher (Haematopus ostralegus palliatus) eggshells from Texas and the Atlantic coast of the United
States.
Key Words: American shorebirds, eggshell thickness, DDT, bioaccumulation, pesticides, chemical pollutants.
Significant eggshell thinning has been documented
in many species of wild birds 1n recent years, nearly all
of it attributable to the effects of p,p’-DDE, a
breakdown metabolite of DDT (Cooke 1973: Stickel
1975: Peakall 1975). Thinning has been most severe in
the eggshells of bird- and fish-eating species (Ratcliffe
1970: Anderson and Hickey 1972), but there have
been few investigations of eggshell thickness in middle
trophic-level birds, including shorebirds of the sub-
order Charadrii. Minor shell thickness changes were
found for Pluvialis apricaria and Tringa nebularia by
Ratcliffe (1970), and for Philohela minor by Dilworth
et al. (1972) and Kreitzer (1972).
Studies of organochlorine residues in shorebirds or
their eggs are more numerous. Keith and Gruchy
(1972) summarized data from several sources, and
additional figures for body and egg residue levels have
been reported for various species by Enderson and
Berger (1968), Flickinger and King (1972), White
et al. (1973), Peakall (1976), and Walker (1977).
Owing to its importance as a game species, several
studies are available on residue levels in American
Woodcocks ( Philohela minor) (Pearce 1971: McLane
et al. 1971, 1973; Dilworth et al. 1972, 1974; Clark and
McLane 1974).
Most of these studies indicated that organochlorine
residues, including DDE, rarely exceeded 4 ug/g wet
weight; however, significantly higher levels of DDE
were found in the bodies of Numenius americanus
(14.0 ug/g wet weight) in Alberta (Peakall 1976),
Philohela minor (13.0 ug/g wet weight) in a heavily
sprayed area in New Brunswick (Pearce 1971),
Pluvialis dominica (62.0 ug/g lipid), and Numenius
phaeopus (39.0 ug/g lipid) in Alaska (Walker 1977),
Vol. 93
LD-NATURALIST
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1979
and in the eggs of Philohela minor (14.9 ug/g wet
weight) in New Brunswick (Dilworth et al. 1972). Ina
DDT-sprayed area around Churchill, Manitoba, the
invertebrate prey of shorebirds contained DDE
residues of 0.3 and 0.4 ug/g wet weight, and shore-
birds there accummulated DDE residues of up to an
average of 39.4 ug/g wet weight (Brown and Brown
1970).
The DDE residue levels reported in these latter
studies were at least as high as those associated with
serious eggshell thinning in certain raptors and fish-
eating birds (Blus et al. 1974; Peakall 1976; Kiff et al.,
in press), yet there have been no reports of such
changes in the eggshells of these or other shorebirds.
This study was undertaken to determine whether
significant changes in eggshell thickness of American
shorebirds have occurred since the introduction of
DDT in the mid-1940s.
Methods
Empty dry eggshells of charadriine shorebirds in
the collection of the Western Foundation of Verte-
brate Zoology were weighed to the nearest 0.001 g on
a Mettler P120 balance, and their length and breadth
were measured to the nearest 0.01 mm with Helios
dial calipers. A shell thickness index (shell
weight X 100/length X breadth) was calculated for
each eggshell; such an index is correlated with actual
eggshell thickness (Anderson and Hickey 1972). Eggs
that were broken, that had blowholes greater than
3mm in diameter, or that were collected in an
advanced stage of incubation, were excluded from the
analyses. The nomenclature and species sequence
used here follows Morony et al. (1975).
Results
The mean thickness indices for pre-1947 (before-
DDT) and post-1947 (since-DDT) North American
shorebird eggshells are shown in Table |. Seven
species had slightly thicker eggshells in the recent
samples, four showed no change, and 16 species had
thinner eggshells in the post-1947 samples. Eggs of
NOTES 189
another species, Charadrius wilsonia, were thicker in
Florida but thinner in Texas than pre-1947 indices.
The maximum amount of difference between the pre-
and post-1947 samples were -4.7% in the “American”
Oystercatcher (Haematopus ostralegus palliatus) in
Texas and along the southern Atlantic coast of the
United States. Recent eggshells of the “black”
Oystercatcher (H. o. bachmani) in California and
Oregon were 3.3% thinner than the mean thickness of
the pre-1947 sample.
Many shorebirds included in this study migrate and
winter in South American countries where DDT is
still intensively used. For comparative purposes, we
also measured eggshells of three resident species of
Chilean shorebirds of three different families
(Table 2). As with the North American species, only
minor changes were noted. The only statistically
significant difference, -4.1%, was found in a plover,
Vanellus chilensis.
Discussion
Eggshell thinning exceeding 20% has generally
resulted in reproductive failure and population
declines in the species involved (Keith and Gruchy
1972: Stickel 1975), but the biological significance of
thinning less than 10% is not well understood (Faber
and Hickey 1973). In this study, the maximum
increase (+3.8%) was similar to the maximum decrease
(4.7%) in thickness index. Although statistically
significant, these minor index changes are probably
due to sampling artifacts (e.g., observer error,
insufficient sample size, geographical variation),
rather than pesticide effects. We know of no biological
phenomena which could readily explain an increasing
shell thickness within the span of time these eggs were
collected.
Disruptions in eggshell ultrastructure and chemical
composition that reduced egg hatchability in a
population of Common Terns (Sterna hirundo) were -
attributed to DDE contamination even in the absence
of marked shell thinning (Fox 1976). Thus, the lack of
substantial thinning of American shorebird eggshells
TABLE 2 — Eggshell thickness of Chilean shorebirds
Early Recent
period period No. of eggs® Mean index + SE %
Species (E) (R) E R E R Change
Nycticryphes semicollaris 1934 — 1938 28(15) 0.93 + 0.011
1962 — 1969 19(11) OLS ae OOS = 3
Vanellus chilensis 1934 — 194] 24(8) 1.21 0.017
1960 — 1968 33(10) 1.16+0.011 4.1*
Gallinago paraguaiae 1933 — 1940 40(20) 0.87 + 0.008
1957 — 1969 22(12) 0.88 £ 0.013 + 1.1
“Number of clutches given in parentheses.
*P< 0.05.
190
in recent years does not prove that these species are
free of pesticide-induced reproductive problems.
It is possible that shorebirds have a lower sensitivity
to DDE-induced eggshell thinning than many higher
trophic-level species, in addition to usually possessing
lower residue burdens. Peakall (1975) categorized the
charadriiforms as being “moderately sensitive” to
DDE, based on data on Herring Gulls (Larus
argentatus) presented in Hickey and Anderson (1968).
Although DDE residues in these species have
evidently not reached a level at which they cause
eggshell thinning, migrant shorebirds may still repre-
sent the most important source of DDE contamina-
tion for Arctic raptors, including Peregrine Falcons
(Falco peregrinus) and Gyrfalcons (F. rusticolus).
Several studies have indicated that migratory shore-
birds possess the highest organochlorine residues of
any prey item taken by these falcons (Cade et al. 1968:
Enderson and Berger 1968: White et al. 1973: Walker
1977).
Acknowledgments
We appreciate the support of Ed N. Harrison and
the Western Foundation of Vertebrate Zoology.
Critical review by A. J. Erskine, J. J. Hickey, and
D. B. Peakall enhanced the quality of the paper. Julie
Kiff and Dana Gardner assisted in preparation of the
manuscript.
Literature Cited
Anderson, D. W. and J. J. Hickey. 1972. Eggshell changes
in certain North American birds. Proceedings of the XV
International Ornithological Congress. pp. 514-540.
Blus, L.J., B.S. Neely, Jr.. A. A. Belisle, and R.M.
Prouty. 1974. Organochlorine residues in Brown Pelican
eggs: relation to reproductive success. Environmental
Pollution 7: 81-91.
Brown, J. J. and A. W. A. Brown. 1970. Biological fate of
DDT in a sub-arctic environment. Journal of Wildlife
Management 34: 929-940.
Cade, T. J., C. M. White, and J. R. Haugh. 1968. Pere-
grines and pesticides in Alaska. Condor 70: 170-178.
Clark, D.R., Jr. and M. A.R. McLane. 1974. Chlori-
nated hydrocarbon and mercury residues in woodcock in
the United States, 1970-1971. Pesticides Monitoring
Journal 8: 15-22.
Cooke, A. S. 1973. Shell thinning in avian eggs by environ-
mental pollutants. Environmental Pollution 4: 85-52.
Dilworth, T.G., J. A. Keith, P. A. Pearce, and L.M.
Reynolds. 1972. DDE and eggshell thickness in New
Brunswick woodcock. Journal of Wildlife Management
36: 1186-1193.
Dilworth, T.G., P. A. Pearce, and J. V. Dobell. 1974.
DDT in New Brunswick woodcocks. Journal of Wildlife
Management 38: 331-337.
Enderson, J.H. and D.D. Berger. 1968. Chlorinated
hydrocarbon residues in peregrines and their prey species
from northern Canada. Condor. 70: 149-153.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Faber, R.A. and J.J. Hickey. 1973. Eggshell thinning,
chlorinated hydrocarbons, and mercury in inland aquatic
bird eggs, 1969 and 1970. Pesticides Monitoring Journal
7: 27-36.
Flickinger, E.L. and K. A. King. 1972. Some effects of
aldrin-treated rice on Gulf Coast wildlife. Journal of
Wildlife Management 36: 706-727.
Fox, G. A. 1976. Eggshell quality: Its ecological and
physiological significance in a DDE-contaminated Com-
mon Tern population. Wilson Bulletin 88: 459-477.
Hickey, J.J. and D.W. Anderson. 1968. Chlorinated
hydrocarbons and eggshell changes in raptorial and fish-
eating birds. Science 162: 271-273.
Keith, J. A. and I. M. Gruchy. 1972. Residue levels of
chemical pollutants in North American birdlife. Pro-
ceedings of the XV International Ornithological Congress.
pp. 437-454.
Kiff, L. F., D. B. Peakall, and S. R. Wilbur. Recent changes
in California Condor eggshells. Condor. /n press.
Kreitzer, J. F. 1972. Thickness of the American Woodcock
eggshell, 1971. Bulletin of Environmental Contamination
and Toxicology 9: 281-286.
McLane, M.A.R., L.F. Stickel, and J.D. Newsom.
1971. Organochlorine pesticide residues in woodcock,
soils, and earthworms in Louisiana, 1965. Pesticides
Monitoring Journal 5: 248-250.
McLane, M. A.R., L. F. Stickel, E. R. Clark, and D.L.
Hughes. 1973. Organochlorine residues in woodcock
wings, Il states — 1970-71. Pesticides Monitoring
Journal 7: 100-103.
Morony, J.J., Jr., W.J. Bock, and J. Farrand, Jr.
1975. Reference list of birds of the world. American
Museum of Natural History, New York. 207 pp.
Peakall, D. B. 1975. Physiological effects of chlorinated
hydrocarbons on avian species. /m Environmental
dynamics of pesticides. Edited by R. Haque and V. H.
Freed. Plenum Publishing Company, New York. pp.
343-360.
Peakall, D.B. 1976. The peregrine falcon (Falco pere-
grinus) and pesticides. Canadian Field-Naturalist 90:
301-307.
Pearce, P. A. 1971. Side effects of forest spraying in New
Brunswick. Transactions of the Thirty-sixth North Ameri-
can Wildlife and Natural Resources Conference. pp.
163-170.
Ratcliffe, D. A. 1970. Changes attributable to pesticides in
egg breakage frequency and eggshell thickness in some
British birds. Journal of Applied Ecology 7: 67-115.
Stickel, W. H. 1975. Some effects of pollutants in terrestial
ecosystems. /n Ecological toxicology research. Edited by
A.D. McIntyre and C.F. Mills. Plenum Publishing
Company, New York. pp. 25-74.
Walker, Wayman. 1977. Chlorinated hydrocarbon pol-
lutants in Alaskan Gryfalcons and their prey. Auk
94: 442 447.
White, C.M., W.B. Emison, and F.S.L. Williamson.
1973. DDE in a resident Aleutian Island peregrine
population. Condor 75: 306-311.
Received 2 October 1978
Accepted 2 January 1979
1979 NOTES 19]
Response of Wintering Moose to Mechanical Habitat Rehabilitation
in Alaska
M. SIGMAN
Alaska Cooperative Wildlife Research Unit, University of Alaska, Fairbanks, Alaska 99701
Present address: Ester, Alaska 99725
Sigman, M. 1979. Response.of wintering Moose to mechanical habitat rehabilitation in Alaska. Canadian Field-Naturalist
93(2): 191-193.
Moose (Alces alces gigas) formed winter concentrations following mechanical crushing of vegetation in a 28-yr-old burn.
Compared to a control area, the density of Moose and calf survival in the rehabilitated area were consistently higher in
January—March 1975. Persistent large aggregations, although dynamic in nature, were related to the distribution of crushed
mature hardwood stands because Moose fed on aspen bark. Observed differences between the two areas resulted from the
provision of a concentrated, available, and nutritious food supply in combination with less severe snow conditions.
Key Words: Moose, Alces alces gigas, habitat, rehabilitation, aggregation, behavior.
In 1975, as part of efforts to manage habitat to
stabilize a declining Moose population, Kenai Nation-
al Moose Range (KNMR) staff used LeTourneau tree
crushers to rehabilitate a portion of an area burned in
1947. An important result of the winter crushing of
mature and regrowth vegetation was the immediate
presence of large concentrations of Moose in the
crushed area. My objectives in observing these
concentrations were to compare the use of the area to
that of a control area and to observe behavioral
responses of Moose to the habitat disturbance.
Study Area
The Willow Lakes Rehabilitation Area (WLRA) is
located in the KNMR in the northwestern portion of
the Kenai Peninsula, Alaska, and is part of a large
plain composed of flats, low ridges, hillocks, muskeg,
and numerous small lakes. The particular area
selected to be crushed could be easily reached froma
road and the vegetation was considered representative
of the 1947 burn. A 460-ha (1137-acre) doughnut-
shaped area was crushed while the “doughnut hole”
was scheduled for a later controlled burn.
Two areas were surveyed by use of fixed-wing
aircraft: a 2330-ha (5750-acre) area which included
crushed and uncrushed portions and a control area of
similar size approximately 6 km away in the 1947
burn. Both areas included many lakes; the actual area
used by Moose for feeding and bedding was roughly
equivalent.
J. L. Oldemeyer, Denver Wildlife Research Center
(unpublished data) has provided detailed information
on the vegetation in the project area before and after
crushing. In general, before crushing, islands of
remnant mature White Spruce (Picea glauca)/
Trembling Aspen (Populus tremuloides)/ Paper Birch
(Betula papyrifera) stands were interspersed with
regrowth birch-spruce (P. glauca and P. mariana)
stands.
Methods
Semi-monthly surveys of the WLRA and control
area were flown from 2 December 1974 until 2 May
1975. I used information from these surveys to select
suitable areas for viewing large concentrations of
Moose. I conducted long (4-8 h) observations of large
aggregations in the crushed area and noted all Moose
in the general area on my way to and from specific off-
road observation sites of aggregations. In addition, I
conducted a daily road survey along the southern
boundary of the project area. Owing to lack of access-
ibility, however, I made no attempt to observe Moose
in the control area. Ground observations continued,
weather permitting, from 14 February until 6 April
ISHS.
During long observation periods, I observed aggre-
gations from natural blinds (e.g., a clump of birch
trees on a nearby hill) approximately 200 m from the
Moose. After allowing a period for the animals to
resume undisturbed behavior after my arrival at the
blind, I recorded data at 15-min intervals, noting the
number of Moose visible from my vantage point, the
number of Moose in aggregations, the general
behavior of individual Moose (e.g., feeding vs. lying),
and the presence of collared Moose (collared previ-
ously by the Alaska Department of Fish and Game). I
also sketched the relative locations of individuals.
Between these observations, I scanned the area
frequently for movements, arrivals and departures in
aggregations, and interactions.
Results
The WLRA received much more use than the
control area. Aerial observations documented 1374 d
of Moose use in the WLRA compared to 466 d in the
control area. Observed densities ranged from 2.5 to
9.8 Moose per km? in the WLRA compared to 1.3 to
3.1 Moose per km? in the control area (Table |). The
higher densities in the crushed area built up between
192
TABLE 1—Comparison of Moose densities observed in the
Willow Lakes Rehabilitation Area (WLRA) and the
control area
Moose/ km2
Date of survey WLRA Control
23 Dec./74 25)
24 Dec./74 3
10 Jan./75 6.3 ral
20 Jan./75 7.0 De
SEkeby 5 6.3 2.0
20 Feb./75 9.8 2.6
6 Mar./75 VD 1.6
14 Mar./75 6.2 DM
1 Apr./75 7.8 2.4
18 Apr./75 4.2 3
early and mid-winter, then decreased in late winter,
while the densities in the control area remained low
throughout the winter.
Calf survival was much higher in the WLRA (Table
2). Cow-calf ratios ranged from 26 to 62 calves per 100
adults, compared with those in the control area, which
decreased steadily from 36 calves per 100 adults to
none by 18 April.
Ground observations in the WLRA documented an
additional 324 d of use by Moose. Although group-
ings of one, two, or three Moose were commonly
observed travelling or feeding in any portion of the
area, all larger groups were observed in crushed
mature hardwood stands. The animals initially con-
sumed branches and twigs of birch and aspen trees.
Concentrations persisted in these area for several days
apparently to consume aspen bark from the downed
mature trees. Based on a vegetation type map of the
TABLE 2—Comparison of calves per 100 adults observed in
the Willow Lakes Rehabilitation Area (WLRA) and the
control area
Calves/ 100 adults
Date of survey WLRA Control
23 Dec./74 39
24 Dec./74 36
10 Jan./75 2 23
20 Jan./75 31 28
5 Feb./75 34 13
20 Feb./75 28 13
6 Mar./75 ; 37/ 24
14 Mar./75 27 II
1 Apr./75 31 4
18 Apr./75 62 0
THE CANADIAN FIELD-NATURALIST
Vol. 93
area, maximum densities within these crushed stands
varied from 0.25 to 30 Moose per km3 (n = 34).
Concentrations persisted but the aggregations
themselves were extremely dynamic during the long
observation periods. Repeated observations of collar-
ed Moose indicate that the individual length of stay
was variable, ranging from | to 112 d, which suggests
that considerable turnover occurred throughout the
winter.
With the exception of cow-calf pairs, Moose rarely
left or entered the stands in groups. Individuals rarely
interacted even at high densities. Only three brief
interactions occurred during 48.1 h of observation
involving 405 Moose.
Discussion
The comparison of Moose densities within the
WLRA and those in the control area demonstrates the
rapid build-up of Moose following crushing and a
prolonged use of the crushed area by many Moose. The
observed mid-winter aggregations, although not
notably larger than those observed in late-winter
situations (Timofeeva 1967; Peek et al. 1974; Rose-
neau and Stern 1974), were extremely dense within the
crushed mature hardwood stands.
The lowland winter range of the 1947 burn is
generally considered poor winter range (Oldemeyer et
al. 1978). Thus, it is likely that the major factor
responsible for the aggregations and concentrations
of Moose in the WLRA was a food supply that was
highly nutritious (J. L. Oldemeyer, personal com-
munication), concentrated, and available.
Snow conditions were also variable between the
two areas. Movement of the large crushers to various
sites compacted the snow into trails and the removal
of vegetation resulted in wind compaction of newly-
fallen snow. Sigman (1977) noted that the increasing
snow depths in typical 1947 burn habitats, in the
nearby Moose Research Center, appeared related to
the timing of calf deaths. In the WLRA, the reduction
of energy expenditure necessary to move through
deep snow and to locate food was combined with an
increased energy intake from the food supply. This
combination may have acted to reverse some of the
stressful effects of typical winter conditions and
seemed to have benefitted calves in particular.
The immediate and prolonged use of the rehabili-
tated area demonstrates flexible aspects of Moose
behavior patterns. Use of the disturbed area confirms
the ability of individual Moose opportunistically to
locate a small area where winter conditions are more
favorable for survival and the ability of wintering
Moose to tolerate high densities.
Acknowledgments
Funding for this study was provided through
Pittman-Robertson funds, administered by the
1979
Alaska Department of Fish and Game (ADFG) and
the Alaska Cooperative Wildlife Research Unit.
Logistical support was provided by the staff of the
Kenai National Moose Range and the Moose Re-
search Center, Kenai, Alaska. In particular, Robert
Ritchey, KNMR, provided aerial sightings and Paul
Arneson, ADFG, provided some ground sightings.
John Oldemeyer, KNMR, and Lyman Nichols,
ADFG, reviewed the manuscript and provided help-
ful suggestions.
Literature Cited
Oldemeyer, J. L., A. W. Franzmann, A. L. Brundage, P. D.
Arneson, and A. Flynn. 1978. Browse quality and the
Kenai Moose population. Journal of Wildlife Manage-
ment 41(3): 533-542.
NOTES 193
Peek, J. M., R. E. LeResche, and D. R. Stevens. 1974. Dy-
namics of Moose aggregations in Alaska, Minnesota, and
Montana. Journal of Mammalogy 55(1): 126-137.
Roseneau, D. G. and P.M. Stern. 1974. Distribution of
Moose, Muskox, and Sheep in northeastern Alaska,
1972. In Distribution of moose, sheep, muskox, and fur-
bearing mammals in Northeastern Alaska. Edited by
R. D. Jakimchuk. Canadian Arctic Gas Study Limited.
Biological Report Series, Volume VI, Chapter I.
Sigman, M. J. 1977. The importance of the cow-calf bond
to overwinter moose calf survival. M.Sc. thesis, University
of Alaska, Fairbanks, Alaska. 185 pp.
Timofeeva, E. K. 1967. Onthe behavior of moose, based on
observations made in the northeastern Leningrad oblast.
Vestnik Leningradskogo Universiteta, Number 15.
(Translation). 11 pp.
Received 2 October 1978
Accepted 3 January 1979
New Localities for the Northern Spring Salamander and the
Four-toed Salamander in Southwestern Quebec
DAVID M. GORDON
Box 180, Macdonald College Post Office, Quebec H9X 1C0
Gordon, David M. 1979. New localities for the Northern Spring Salamander and the Four-toed Salamander in south-
western Quebec. Canadian Field-Naturalist 93(2): 193-195.
New localities for the salamanders Gyrinophilus porphyriticus porphyriticus and Hemidactylium scutatum are documented
and the habitat at the collecting sites described. This is the first record for Gyrinophilus west of the Richelieu River and only
the second record outside the Appalachian Mountain region. Hemidactylium is reported for the second time south of the St.
Lawrence River in Quebec.
Key Words: salamanders, Gyrinophilus, Hemidactylium, geographic distribution, Quebec.
The geographic distribution of salamanders in
Quebec is poorly known (see Figure 1). The secretive
nature and seasonally variable habits of these animals
and, until recently, the lack of systematic field work
have contributed to this situation. The recent efforts
of the author in conjunction with the National
Museum of Natural Sciences, and of Weller (1977)
have expanded our knowledge of the nature of the
distributions of. two of the species of salamanders in
Quebec. This note documents two new localities for
the Northern Spring Salamander and one new locality
for the Four-toed Salamander.
Gyrinophilus porphyriticus porphyriticus
During the course of field work (15 August 1973; 29
April and 19 May 1975) two larval and 10 meta-
morphosed Northern Spring Salamanders (Gyrino-
Philus p. porphvriticus) were discovered in the
headwaters of Riviere aux Outardes-Est, near Frank-.
lin Centre, Huntingdon County (45°01’N, 73°54’W),.
Riviére aux Outardes-Est originates at a swamp
and small lake at the top of Covey Hill at an elevation
of 210-225 m. At the collecting site the stream cuts
through a stand of Eastern Hemlock (7suga cana-
densis) and mixed hardwoods of Sugar Maple (Acer
saccharum), American Beech (Fagus grandifolia),
and Yellow Birch (Betula alleghaniensis). The stream
varies in width from 4 to 10 m, with steep banks
ranging from 10 to 80cm in height. The stream
bottom consists of gravel and sand with some detritus;
rocks of all shapes and sizes provide cover in the
stream and along the banks. An extensive network of
passages among the rocks occurs where the stream has
eroded the bank.
The Northern Dusky Salamander ( Desmognathus
fuscus fuscus) occurs in the seepage areas along the
194
banks and the Northern Two-lined Salamander
(Eurycea bislineata bislineata) is abundant through-
out the stream. The Spotted Salamander (Amby-
stoma maculatum), the Blue-spotted Salamander
(Ambystoma laterale), and the Red-backed Sala-
mander (Plethodon cinereus cinereus) occur in the
woods adjacent to the stream.
This is the first record of the Spring Salamander
west of the Richelieu River, and only the second
record of the species from outside of the Appalachian
Mountain region of Quebec. Weller (1977) reported
Spring Salamanders from Yamaska Mountain of the
Monteregian Hills on the St. Lawrence Lowlands.
More recently, he has taken four metamorphosed
Gyrinophilus from one locality on Shefford Moun-
tain (Figure 1) at the edge of the Appalachian
Mountain range. The Dusky Salamander and the
Two-lined Salamander were also found at this
locality. These specimens and those I collected have
76° 75°
THE CANADIAN FIELD-NATURALIST
Vol. 93
been deposited in the herpetological collection of the
National Museum of Natural Sciences as NMC
17816, and NMC 16379, 16767, 16791 respectively.
Hemidactylium scutatum
The Four-toed Salamander (Hemidactyvlium scu-
tatum) has been reported from four scattered locali-
ties in Quebec of which only one is south of the St.
Lawrence River (Figure 1). On 29 April 1976, two
metamorphosed Hemidactylium were taken from a
site 11 km E of Covey Hill, Huntingdon County,
Quebec (45°01’N, 73°38’W).
The collecting area was a woodlot of Eastern
Hemlock, Sugar Maple, American Beech, and Yellow
Birch. The salamanders were discovered in a pile of
bark at the base of a dead tree, close to a small
woodland pond. Moss covering the rocks and logs,
and bark and leaf litter provided cover. The ground
was extremely wet with many scattered pools of
FIGURE |. Solid circles depict localities for Gyrinophilus p. porphyriticus in Canada, triangles the known localities of
Hemidactvlium scutatum in Quebec. Locality 1, Franklin Centre Gyrinophilus record, 2, Shefford Mountain
Gyrinophilus record; 3, the new record for Hemidactylium. The inset maps illustrate the North American ranges
according to Conant (1975): A, Gyrinophilus porphyriticus and B, Hemidactylium scutatum.
1979
temporary water. Sphagnum moss was present only in
small widely scattered patches. This habitat is
virtually identical to an area on Ile Perrot, Quebec
where I have collected Four-toed Salamanders in past
years, and to another Ile Perrot locality reported by
McCoy and Durden (1965). Denman(1961, 1965) and
Gorham (1955) found Hemidactylium in similar
forest types. The Red-back Salamander and the Blue-
spotted Salamander occurred in the immediate
vicinity, and the Red-spotted Newt (Notophthalmus
viridescens) was abundant in the nearby woodland
pond.
This is the second record of Hemidactylium south
of the St. Lawrence River in Quebec. The specimens
are catalogued as NMC 17539.
My thanks go to Barbara L. Brown for her
assistance in the field, to Wayne F. Weller who
provided me with a copy of his manuscript on stream
salamanders in advance of publication, and allowed
me to publish his Gyrinophilus record, and to Francis
R. Cook for his advice and encouragement regarding
the field work and the manuscript. Portions of this
study were financed by the National Museum of
NOTES 195
Natural Sciences, as part of a herpetofaunal survey in
Quebec in 1975.
Literature Cited
Conant, R. 1975. A field guide to reptiles and amphibians
of eastern and central North America. 2nd edition.
Houghton Mifflin Company, Boston. xvii + 429 pp.
Denman, N.S. 1961. A range extension of the Four-toed
Salamander in eastern Canada. Canadian Field-Natur-
alist 75: 110.
Denman, N.S. 1965. Further records of the Four-toed
Salamander with remarks on its habitat in Quebec
province. Canadian Field-Naturalist 79: 76-77.
Gorham, S. W. 1955. Notes on the Four-toed Salamander
in the province of Quebec. Canadian Field-Naturalist
69: 167.
McCoy, C.J. and C.J. Durden. 1965. New distribution
records of amphibians and reptiles in eastern Canada.
Canadian Field-Naturalist 79: 156-157.
Weller, W. F. 1977. Distribution of stream salamanders in
southwestern Quebec. Canadian Field-Naturalist 91; 299-
303.
Received 8 April 1978
Accepted I] January 1979
First Record of the Long-tailed Shrew (Sorex dispar)
in New Brunswick
GORDON L. KIRKLAND, JR.,! DAVID F. SCHMIDT,2 and CAROL J. KIRKLAND!
'Vertebrate Museum, Shippensburg State College, Shippensburg, Pennsylvania 17257
?Division of Mammals, National Museum of Natural History, Washington, D.C. 20560
Kirkland, Gordon L., Jr., David F. Schmidt, and Carol J. Kirkland. 1979. First record of the Long-tailed Shrew (Sorex
dispar) in New Brunswick. Canadian Field-Naturalist 93(2): 195-198.
One subadult male Long-tailed Shrew (Sorex dispar) was collected in Albert County, New Brunswick, on 25 July 1978. This is
the first New Brunswick record for this species and the fifth Canadian specimen collected. The New Brunswick specimen was
compared to Sorex gaspensis and New England S. dispar to ascertain its specific identity.
Key Words: Sorex dispar, Long-tailed Shrew, New Brunswick, first record, Sorex gaspensis.
The Long-tailed Shrew (Sorex dispar) is known
from Canada on the basis of four specimens (Ameri-
can Museum of Natural History 17434447) collected
at two localities in Quebec near the United States
border: 16 km (10 mi) S of Armstrong, near Lac du
Portage; and south of Cartierville, just a few yards
north of the New Hampshire border (Peterson 1966)
(Figure 1). With the exception of these specimens
which at present cannot be located for study, S. dispar
is confined to the United States in the Appalachians
and adjacent mountains in a narrow belt extending
from Maine to North Carolina (Kirkland and Van
Deusen 1979). The closely related Gaspé Shrew
(Sorex gaspensis) is an endemic Canadian species
which until recently was thought to be restricted to the
Gaspé region of Quebec. But it has recently been
collected in such disjunct locations as Mount Car-
leton, New Brunswick (Peterson and Symansky 1963)
and Cape Breton Island, Nova Scotia (Roscoe and
Majka 1976) (Figure 1). In an attempt to determine
whether the hiatus in the distribution of S. gaspensis is
real or the product of inadequate sampling, field work
was conducted in southeastern New Brunswick and in
the Cobequid Mountains and on North Mountain in
196 THE CANADIAN FIELD-NATURALIST
GASP E
fo}
NEW
BRYN SW I
FIGURE |.
Vol. 93
PEON SOLA
g
fr
sS
PRINCE EDWARD
ISAM CAPE BRETON
ISLAND
Kilometers
Location of capture sites of Sorex gaspensis (open circles), Maine Sorex dispar (dots), SSC 8393 (dot within
circle), and previously recorded Quebec S. dispar (triangles), based on Godin (1977), Kirkland and Van Deusen (1979),
and Peterson (1966). One mark may represent more than one locality if several specimens have been collected from
different localities in the same region.
Nova Scotia during July 1978.
In 2057 trapnights (TN) of sampling effort at six
localities in Nova Scotia and New Brunswick, no S.
gaspensis were collected; however, on 25 July 1978, a
single subadult male S. dispar was trapped 5.3 km N,
3.5 km W of Riverside-Albert, Albert County, New
Brunswick. This locality is approximately 305 km
ESE of the nearest previous locality for this species at
Beaver Creek, Aroostook County, Maine (Godin
1977). The New Brunswick specimen (Shippensburg
State College 8393) was captured in a Museum
Special trap baited with rolled oats and set in rocks,
30cm below the surface, on a rocky east-facing
hillside in a deciduous-coniferous forest (elevation
180 m). The capture site was dominated by Yellow
Birch (Betula alleghaniensis) with Mountain Maple
(Acer spicatum) and Balsam Fir (Abies balsamea) as
subdominants. Living ground cover at the site
averaged 51-75% and consisted of mosses, ferns, and
seedlings of Mountain Maple. In 306 TN of sampling
effort, 23 small mammals representing six species
were captured in the trapline that yielded the single S.
dispar: seven Smoky Shrews (Sorex fumeus), four
Short-tailed Shrews ( Blarina brevicauda), one North-
ern Flying Squirrel (Glaucomys sabrinus), eight
Southern Red-backed Voles (Clethrionomys gap-
peri), and two Woodland Jumping Mice (Napaeo-
zapus insignis). The trap in which the S. dispar was
captured had yielded a male S. fumeus the previous
morning (24 July). The standard external measure-
ments and 16 selected cranial and mandibular
characters for specimen SSC 8393 are presented in
Table |.
Because the New Brunswick specimen was captured
near the center of the disjunction in the range of S.
gaspensis and over 300 km from the nearest S. dispar
1979
TABLE !—Measurements for six external and 16 cranial/
mandibular characters of New Brunswick specimen
SSC 8393 and the ranges of measurements for these 22
characters in 23 S. gaspensis and 56 New England S. dispar
(All linear measurements in millimetres; weight in grams)
Character SSC Range
Does S. gaspensis S. dispar
Total length 123 95-127 103-136.5
Tail length 56 45- 55* 46- 61
Hind foot length 13 10.5 -12.5* 12- 15
Ear length 8 5- 9 5- 9
Body length 67 45-77 48- 79
Weight 4.2 2.2 - 4.3 4.0 - 4.9
Greatest length [ASS ISAS les* svleS a 18e3,
Condylobasal length 16.8 15.35-16.35* 16.45-17.7
Interorbital breadth 3.15 2.8 — 3.5 3.1 — 3.6
Cranial breadth 8.25 Toll = Sa 7.5 — 8.3
Molariform tooth
TOW 3.9 Bre 3-69%80 3/5 — 43
Cheek tooth row 4.55 4.05-— 4.4* 4.1 - 49
Total tooth row 7.05 6.35- 7.25 6.8 — 7.6
Incisor width 1.15 1.0 - 1.3 1.1 - 1.4
Canine width 1.65 ES lRS Sth elE4S eS:
Molar width 3.8 3.2 — 3.8 3.65— 4.15
Nasal length 6.25 5.4 — 6.3 5.6 — 7.25
Palatal length V2 6.0 — 6.75* 6.6 — 7.5
Post-palatal length 8.2 6.9 — 7.6* 7.3 — 8.35
Mandible length I 10.15 9.1 -— 9.9* 9.75-10.75
Mandible length II —:11.00 9.7 -10.8* 10.1 -11.35
Mandible height 32 2.95— 3.95 3.05— 4.0
*SSC 8393 exceeds range for character.
locality, it was important to ascertain its specific
identity correctly. Was it a S. dispar, a S. gaspensis, or
was it intermediate between these two taxa? The
existence of an intergrade specimen at this New
Brunswick locality would call into question the
specific status of S. gaspensis (Kirkland and Van
Deusen 1979).
Univariate and multivariate statistical analyses
were performed to compare the New Brunswick
specimen with 23 S. gaspensis and 56 New England S.
dispar. For localities of these specimens and
descriptions of the measurements see Kirkland and
Van Deusen (1979). All skulls were measured by the
senior author with Helios dial micrometers (calibra-
tion 0.05 mm) under a dissecting microscope.
Comparisons of the New Brunswick specimen with
S. gaspensis and New England S. dispar revealed that
the New Brunswick specimen exceeded the size range
of S. gaspensis for 12 of 22 characters and equalled the
largest known specimen for one other (Table 1). The
New Brunswick specimen fell within the size range of
New England S. dispar for all 22 characters (Table 1).
Data from 247 S. dispar with locality latitudes
ranging from from 35.6° N (North Carolina) to
NOTES
OF
45.9° N (Maine) were analyzed with the SPSS
Computer Package (Nie et al. 1975) to produce
regression lines of size versus latitude for each of 22
characters. These regression equations were used to
predict the characteristics of a S. dispar from the
latitude of the New Brunswick specimen (45.8° N).
The New Brunswick specimen fell within the 95%
confidence interval of the predicted value (see Zar
1974) for 21 of 22 characters (Table 2). In addition,
discriminant function analyses (BMD 07M, Dixon
1967) were performed on the New Brunswick speci-
men, 14 S. gaspensis, and 25 New England S. dispar
using 18 characters. The New Brunswick specimen
was assigned by the computer analysis to S. dispar
both when assigned to S. gaspensis ( P = 0.753) and to
New England S. dispar (P= 1.000). The respective
discriminant scores (first two canonical variables) for
SSC 8393, the 14 S. gaspensis, and 25 S. dispar were
= 0358s = L010 x = 2 965. \(range —45I 9 ato
=12395)), Yi = —0:000) (range: = —1-956n tom 24 Nl):
X= 1779 (range 01169) to 431479), Yo= 0.000
@Gange™ —1-499) to 12788), when) SSG 8393
was assigned a priori to S§. gaspensis, and
X = +3.415, Y=-0.010; X= -4.402 (range -5.944
TABLE 2—Comparisons of the observed values of 22
morphological variables in New Brunswick Long-tailed
Shrew (Sorex dispar) specimen (SSC 8393) from 45.8°N
with the predicted values and 95% confidence intervals of a
Sorex dispar from that latitude based on regression analysis
of 247 S. dispar collected at latitudes 35.6°-45.9°N
95%
Character SSC _ Predicted Confidence
8393 value interval
Total length 123 115.92 103.76-128.08
Tail length 56 54.68 49.02- 60.34
Hind foot length 13 13.06 11.51- 14.61
Ear length 8 7.54 4.51- 10.57 ©
Body length 67 61.35 5056= 72514
Weight 4.2 4.37 DM= O03}
Greatest length 17.85 18.02 17.34— 18.70
Condylobasal length 16.8 16.97 16.23- 17.71
Interorbital breadth 3.15 3.47 3.13— 3.81
Cranial breadth 8.25 7.87 V3= B37
Molariform tooth row 3.90 4.07 3.87— 4.27
Cheek tooth row 4.55 4.46 4.22— 4.70
Total tooth row 7.05 7.30 6.98=— 7262
Incisor width 1.15 1.36 1.22— - 1.50
Canine width 1.65 1.51 les38= 1k69
Molar width 3.80 3.89 36l= 4217
Nasal length 6.25 6.20 5.62— 6.78
Palatal length V2) 7.04 6.64— 7.44
Post-palatal length 8.20 7.94 7.46— 8.42
Mandible length I 10.15 10.08 9.66— 10.50
Mandible length II 11.00 11.01 10.49— 11.53
Mandible height 3.20 3.50 3.12— 3.88
198
to —2.916), Y =-0.000 (range -1.708 to +1.595);
XK = +2.371 (range -0.633 to +4.744), Y = -0.000
(range -2.226 to +1.901) when SSC 8393 was
assigned a priori to S. dispar.
The results of these three analyses suggest that the
New Brunswick specimen is a Sorex dispar and
resembles S. dispar from New England.
Acknowledgments
The field work in Nova Scotia and New Brunswick
was supported by a grant from The Explorers Club.
We thank Henry W. Setzer and personnel of the New
Brunswick Department of Natural Resources and the
Nova Scotia Department of Lands and Forests for
their assistance. We acknowledge Don E. Wilson and
Michael A. Bogan for critically reviewing this
manuscript.
Literature Cited
Dixon, W.J. 1967. BMD Biomedical Computer Pro-
grams. University of California Press, University of
California Publications in Automatic Computation,
Number 2. 600 pp.
Godin, A. J. 1977. Wild mammals of New England. Johns
THE CANADIAN FIELD-NATURALIST
Vol. 93
Hopkins University Press, Baltimore. 304 pp.
Kirkland, G. L., Jr. and H. M. Van Deusen. 1979. The
shrews of the Sorex dispar group: Sorex dispar Batchelder
and Sorex gaspensis Anthony and Goodwin. American
Museum of Natural History Novitates. Jn press.
Nie, N. H., C. H. Hull, J. G. Jenkins, K. Steinbrenner, and
D.H. Bent. 1975. Statistical package for the social
sciences. 2nd edition. McGraw-Hill Book Company, New
York. 675 pp.
Peterson, R. L. 1966. The mammals of eastern Canada.
Oxford University Press, Toronto. 465 pp.
Peterson, R.S.and A. Symansky. 1963. First record of the
Gaspé Shrew from New Brunswick. Journal of Mam-
malogy 44: 278-279.
Roscoe, B. and C. Majka. 1976. First records of the Rock
Vole (Microtus chrotorrhinus) and the Gaspé Shrew
(Sorex gaspensis) from Nova Scotia and a second record
of the Thompson’s Pygmy Shrew ( Microsorex thompsoni)
from Cape Breton Island. Canadian Field-Naturalist
90: 497-498.
Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall, Inc.,
Englewood Cliffs, New Jersey. 620 pp.
Received 20 November 1978
Accepted 11 January 1979
Intraspecific Food Theft by the American Kestrel
PETER M. FETTEROLF
Department of Zoology, University of Toronto, Toronto, Ontario M5S 1A1
Fetterolf, P. M. 1979. Intraspecific food theft by the American Kestrel. Canadian Field-Naturalist 93(2): 198.
One male American Kestrel, Falco sparverius, was observed in an unsuccessful attempt to steal prey from another male
American Kestrel.
Key Words: American Kestrel, prey theft, Falco sparverius.
On 24 March 1978, I watched two male American
Kestrels, Falco sparverius, at the southern edge of the
Toronto Island Airport, Toronto, Ontario. The
weather was cold (+3°C) and clear. The birds, perched
approximately 5 m apart in 15- to 20-m willows, Salix
rigida, faced the open grassland to the north. At 12:02
EST, one kestrel plunged from its perch and captured
a rodent, probably a vole, Microtus sp. The bird did
not mantle its prey, whereupon the second male
swooped down from its perch, grabbed the vole, and
rapidly flew north low across the airport runways. The
victim took flight immediately, climbed above the
robber and dove upon it three times, shrieking the
“klee” call just prior to contact each time. At each
contact, the pursurer apparently dug its talons into the
fleeing bird. On the third assault, the robber dropped
the rodent and departed. The other kestrel pounced
on the prey at once, assumed an exaggerated mantling
posture with the wingtips nearly touching, and ate the
rodent. The theft, chase, and successful recapture of
the prey took less than 90 s and covered a distance of
about 40 m.
No reports of prey theft by American Kestrels were
found but a European Kestrel, F. tinnuculus, robbed
prey from a Short-eared Owl, Asio flammeus (Boyle,
G. L. 1974. Kestrel taking prey from Short-eared
Owl. British Birds 67: 474-475). Prey-robbing by
kestrels seems to be rare, but it is not surprising that
the observed encounter occurred at a time of the year
when food supplies were likely to be low.
Special thanks go to Keith Bildstein for comment-
ing on an earlier draft.
Received 3 November 1978
Accepted 14 January 1979
1979 NOTES 199
First Record of the Northern Brook Lamprey, Ichthyomyzon fossor,
in the Nelson River Drainage, Manitoba
J. JYRKKANEN!? and D. G. WRIGHT3
‘Manitoba Department of Renewable Resources, Hadashville, Manitoba
*Present address: Box 396, Garibaldi Highlands, Squamish, British Columbia VON ITO
‘Fisheries and Environment Canada, Fisheries and Oceans, Freshwater Institute, 501 University Crescent, Winnipeg,
Manitoba R3T 2N6
Jyrkkanen, J. and D. G. Wright. 1979. First record of the Northern Brook Lamprey, /chthyomyzon fossor, in the Nelson
River drainage, Manitoba. Canadian Field-Naturalist 93(2): 199-200.
Fourteen specimens of Northern Brook Lamprey, /chthyomyzon fossor, were collected from the Birch River, a tributary to
the Winnipeg River in southeastern Manitoba on 8 and 13 May 1977. This represents the first record of this species in the
Nelson River drainage and is a significant extension of the presently known range.
Key Words: Ichthyomyzon fossor, range extension, Nelson River drainage, Manitoba, new records, geographical
distribution.
Fourteen specimens of Northern Brook Lamprey,
Ichthyomyzon fossor, were captured by hand from
the Birch River, upstream of the town of Prawda,
Manitoba (49° 39’N, 95°48’W) on 8 and 13 May 1977.
The Birch River is a tributary of the Winnipeg River,
in the Nelson River system, flowing into Hudson Bay.
This represents the first record of this species in the
Nelson River drainage system and Is significant in that
it is a range extension of over 500 km from the nearest
known population.
Identification of the specimens was confirmed by
E. J. Crossman of the Royal Ontario Museum.
Morphological characteristics on which species iden-
tification is based are as follows: lateral teeth all uni-
cuspid; the supraoral lamina a single bicuspid tooth;
infraoral lamina cusps blunt; diameter of the sucking
disc less than one half the length of the branchial
region; myotomes in the trunk, between the last gill
opening and the anus, number between 49 and 54;
total length of mature individuals between 100 and
255 mm. The specimens have been deposited in the
collection of the Royal Ontario Museum, Toronto
and are catalogued under the number ROM 34264. Of
the 14 specimens collected, 10 were males, averaging
116.5 mm (SE =3.6 mm), and four were females,
averaging 136.5 mm (SE = 9.6 mm) in total length. All
specimens were sexually mature with females having
free ova within the body cavity. It is assumed that they
were spawning at the time of collection.
Ichthyomyzon fossor is currently known to exist
over a very limited range. Hubbs and Trautman
(1937) report that the species is abundant in all of the
Great Lakes drainages of the State of Michigan (Erie,
Huron, Michigan, and Superior) and from Scott
Creek in the Mississippi River drainage of Wiscon-
sin. Leach (1940) reports the occurrence of the species
in the Tippecanoe River (Mississippi River drainage)
in northern Indiana. Dymond (1947) indicates that J.
fossor was resident in the Thames River system, while
Vladykov (1949) reports that specimens were collect-
ed in the Yamaska and St. Francis rivers of southern
Quebec. Scott and Crossman (1973) add that the
distribution of the species includes Georgian Bay,
Lake Nipissing, and the north shore tributaries of
Lake Superior.
The Birch River is a small, fairly slow-moving river
with an estimated maximum flow of 5.7-8.5 m3/s
(200-300 ft3/s) and a low flow of less than 0.15 m3/s
(5 ft3/s). Winter flows may be augmented by overflow
from the Shoal Lake viaduct entering the upper
reaches of the Birch River at East Braintree. The
substrate of the Birch River is highly varied with silty
sediments in the quieter reaches of the stream, gravel
and cobble riffles, bedrock outcroppings, and several
small waterfalls. A thick mat of algae covers the .
shallow, rocky substrates in the summer.
Hubbs and Trautman (1937) state that /. fossor
lives in creeks and small rivers, apparently avoiding
both small brooks and large rivers. It has never been
recorded in lakes, either small or large. This would
seem to preclude the hypothesis that the Birch River
population has been derived from either Lake
Superior or Wisconsin populations. Vladykov (1949)
reports that ammocoetes of /. fossor are sold as bait
for sport fishing in Quebec. This statement tends to
support a hypothesis that the Birch River population
may have been introduced by an angler who dis-
carded his remaining bait after a fishing trip.
Because the species 1s not parasitic, the presence of
I. fossor in the Nelson River drainage basin should not
present any possibility of endangering or destroying
native fishes.
200
Acknowledgments
The authors acknowledge the assistance of D. P.
Scott, Fisheries and Oceans, Winnipeg, C. C. Lind-
sey, Department of Zoology, University of Manitoba,
Winnipeg, and E. J. Crossman, Curator of Ichthy-
ology and Herpetology, Royal Ontario Museum,
Toronto, in confirming the identification of the
specimens, and thank W. Coder and J. Johnson,
Manitoba Department of Renewable Resources,
Hadashville, for their assistance in the field.
Literature Cited
Dymond, J.R. 1947. A list of freshwater fish of Canada
east of the Rocky Mountains, with keys. Royal Ontario
Museum of Zoology, Miscellaneous Publication Number
1. 55 pp.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Hubbs, C. L. and M. B. Trautman. 1937. A revision of the
lamprey genus /chthvomyzon. Museum of Zoology, Uni-
versity of Michigan, Miscellaneous Publication Number
35. 109 pp.
Leach, W. J. 1940. Occurrence and life history of the
northern brook lamprey, /chthvomyzon fossor, in Indi-
ana. Copeia 1940 (1): 21-34.
Scott, W. B. and E. J. Crossman. 1973. Freshwater fishes
of Canada. Bulletin of the Fisheries Research Board of
Canada 184: 49-51.
Vladykov, V. D. 1949. Quebec lampreys ( Petromyzonidae).
1. List of species and their economic importance. Depart-
ment of Fisheries, Province cf Quebec, Contribution
Number 26. 67 pp.
Received 6 November 1978
Accepted 12 January 1979
Blue Grouse Brood Hen — Black Bear Confrontation
M. G. SULLIVAN
Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9
Sullivan, M. G. 1979. Blue Grouse brood hen — Black Bear confrontation. Canadian Field-Naturalist 93(2): 200.
On 14 July 1977, while engaged in a census of Blue
Grouse (Dendragapus obscurus) near Courtenay,
Vancouver Island, British Columbia, I heard a
banded female grouse clucking loudly (indicative of
brood disturbance) as I approached the area. From
about 75 m away I observed the grouse flying at a
Black Bear (Ursus americanus). Flaring about a metre
from the bear, the hen then landed on the ground
about 20 m away, clucking and rushing about. The
bear ran towards the grouse, whereupon she flew at
the bear and again landed some distance away.
Through these maneuvers, the hen seemed to lead the
bear out of a marshy valley onto a rocky ridge. At this
point, the bear appeared to catch my scent and ambled
out of sight. The hen then returned to the marshy area.
Although no chicks were found, I later discovered that
this hen had a brood at that time. Her behavior was
similar to that displayed towards humans when chicks
are present.
Searching the area, I found several stumps that had
been freshly torn apart by a bear searching for insects.
Ripe Huckleberries (Vaccinium ovalifolium) were
plentiful and are an important food for bears in this
area (unpublished data). Thus, a food shortage was
unlikely at the time.
An interesting addition to this incident is that on31
May 1977, I had found a leg band from an adult male
Blue Grouse in a month-old bear scat in the same
general area. This bird was banded as an adult on 16
June 1975, about 2.4 km from where the band was
found and may have been eaten as carrion.
Received 14 February 1978
Accepted 7 November 1978
News and Comment
Editor’s Report for 1978
The Canadian Field-Naturalist received 149 manu-
scripts for consideration in 1978. This number is
consistent with the 147 submitted in 1976 and the 137
uel OMT
In 1978, 77 papers (38 Articles and 39 Notes)
originally submitted from 1975 to 1978 were publish-
ed in Volume 92 of The Canadian Field-Naturalist.
The breakdown according to subject matter is as
follows: birds, 25; mammals, 23; plants, 17; inverte-
brates, 5; amphibians and reptiles, 4; fishes, 2; and
other, 1. Comparison with the number of manuscripts
published in the two previous years, 124 (34 Articles
and 90 Notes) in 1976 and 101 (30 Articles and 71
Notes) in 1977, shows that the number of Notes
published in 1978 was considerably lower. No
explanation is apparent for this; certainly there have
been no particular delays in the publication process
nor any changes in acceptance criteria.
Occasionally there is a long lapse after a manuscript
has been returned to its authors for revision and
before it is resubmitted and accepted for publication.
In fact, this interval has even exceeded three years!
Therefore, the proportions of accepted papers with
respect to the years in which they were submitted
(number accepted/number received) have increased
over the earlier published figures. The revised propor-
tions are as follows: 1974 — 116/152; 1975 — 122/ 167;
1976 — 90/147; and so far for 1977 — 86/137.
The financial position of The Canadian Field-
Naturalist is currently good because several authors
have paid page charges for all published pages rather
than just the obligatory charges for pages over six.
Therefore, for the first time in several years, it was not
necessary to submit an application for a grant to the
Natural Sciences and Engineering Research Council
(formerly the granting body was the National Re-
search Council of Canada). Furthermore, our printer,
M.O.M. Printing of Ottawa, has tried to keep our
printing costs down while at the same time continuing
to put out a quality product. The financial outlook
then for 1979 is conditionally bright.
LORRAINE C. SMITH
Editor
Proposals Invited from Field Research Investigators
The Center for Field Research, a non-profit
organization established to raise private funds for
field research and to encourage public understanding
of science, is currently accepting proposals for 1980.
Projects are considered on the basis of scientific merit
and their ability to utilize the assistance of motivated
lay volunteers in the field.
In the past seven years, The Center for Field
Research and its affiliate, Earthwatch, have raised
over $1.5 million from interested members of the
public who have also contributed their time and skills
to worthy research in a wide variety of disciplines.
This year, it expects to grant another $500,000 in
support of 66 projects, including the observation of
sea cow distribution and behavior, western Australia;
a study of the ecology and social behavior of the
spotted hyena, Kenya; an archaeological and archi-
201
tectural investigation of Repton Anglo-Saxon
church, England; an anthropological study of return-
ing migrants, Newfoundland; a paleontological sur-
vey and fossil collection of Devonian rocks, Idaho;
and an archaeological study of early Iron Age
settlement and economic systems, West Germany.
Proposal deadlines for 1980 research are: | June
1979 (for work beginning after | December 1979): |
October 1979 (for work beginning after | June 1980):
and 15 January 1980 (for work beginning after |
September 1980). Scholars of all nationalities and
from all disciplines are invited to apply. For appli-
cation guidelines write Nancy Bell Scott, The Center
for Field Research, 10 Juniper Road, Box 127-Q,
Belmont, Massachusetts 02178 (Phone 617-
489-3032).
202
Inland Bird Banding
This journal, formerly Inland Bird Banding News,
published by the Inland Bird Banding Association
and printed by Allen Press, has been drastically
changed by its new Editor. It is now a quarterly
refereed journal and will no longer contain the news
and other columns that it once had. These items will
continue in some form in a newsletter.
The Editor is actively soliciting manuscripts that
deal with bird-banding techniques and the results of
bird-banding studies, especially studies dealing with
birds from middle North America (Canada to the Gulf
Coast). Interested ornithologists and students of birds
who have suitable material for publication (preferably
manuscripts of 10 pages or less in length) should
THE CANADIAN FIELD-NATURALIST
Vol. 93
contact the Editor, Jerome A. Jackson, Department
of Biological Sciences, Mississippi State University,
Mississippi State, Mississippi 39762 (phone 601-
325-5722).
Bluebird Society
A new society, the Bluebird Society of North
America, has been formed. Its address is as fol-
lows: Box 6295, Silver Springs, Maryland 20906.
We thank Colleen Hislop for correcting the page proofs for this issue.
Book Reviews
ZOOLOGY
Where to Find Birds in British Columbia
By David M. Mark. 1978. Kestrel Press, New Westminster.
72 pp., illus. Paper $3.00. (Available by mail from
Kestrel Press, P.O. Box 2054, New Westminster,
British Columbia V3L 5A3.)
This small book (13.5 X 21 cm) gives bird-finding
information for 49 areas (“sites”) in British Columbia
(hereafter B.C.), which are described, region by
region, for eight geographic regions of the province. A
sketch map shows the general locations of birding
areas in each region, but the reader will need a good
road map to use with the book. The introduction (six
pages) includes a helpful discussion of bird dis-
tribution in B.C. as related to the Biotic Areas of
Munro and Cowan (B.C. Provincial Museum Special
Publication 2, 1947). Following this, eight pages are
occupied by a photocopy of the recent (1977), com-
prehensive checklist of B.C. birds issued by the
Provincial Museum, which identifies those species
known to breed in B.C. and those of accidental
occurrence. The main part of the book (pages 19 to 63)
is devoted to area accounts, which vary in length from
five lines to two pages. These are generally up-to-date
and informative, with specific and accurate route
directions, including distances (the author is a
geographer!), and usually with lists of particular bird
species to expect. Finally, there is a seven-page section
on “sought-after species” (those most often sought by
birders), with references to area accounts and some-
times additional comments as well. Besides the
author, 15 other individuals or groups contributed
information for the book.
My only major criticism of this guide is that
coverage is poor for some parts of B.C. (As I was one
of the contributors, | am perhaps indicting myself
for not providing more information!) For example,
Vancouver Island (and the Victoria area in par-
ticular) receives rather scanty treatment considering
its diverse avifauna and the heavy visitation it receives
from out-of-province birders. None of the five
national parks in B.C. is mentioned. On the other
hand, coverage is quite good for the Vancouver area
and most of the southern interior, and the accounts
included for those areas can scarcely be faulted. |
sincerely hope the gaps in coverage will be filled ina
future edition.
All in all, this is a very good bird-finding guide, at
least for a pioneering effort in a province previously
lacking one. It will prove very helpful both to B.C.
birders and to those visiting the province from
elsewhere.
WAYNE C. WEBER
Department of Biological Sciences, Mississippi State Uni-
versity, Mississippi State, Mississippi 39762, USA
The Moths of America North of Mexico, Including Greenland.
Fascicle 22.2, Noctuoidea (in part): Lymantriidae
By D.C. Ferguson. 1978. Classey and the Wedge Entomo-
logical Research Foundation (distributed in North
America by Entomological Reprint Specialists, Los
Angeles). US $48 ($40 by subscription).
This fascicle, often advertized as soon appearing,
finally did. It was eagerly awaited by all who will have
to use it. We have to admit that the Lymantriidae area
very difficult family and many things had to be
elucidated to make publication useful: names, distri-
bution, and general considerations.
This begins with the family name, and everybody
concerned is glad that there were no changes in this as
well as in often used generic names. These have been
not quite stable in the past because of international
faunal and taxonomic complications, like Gynae-
phora, Dasychira, Lymantria, Leucoma, Euproctis.
Appreciable conservatism 1s also shown in specific
names, e.g., Euproctis chrysorrhoea. One may hope
that this final nomenclature finds easy access into our
collections and papers.
One might question whether it was necessary to fill
the literature with all the new “subspecific” names.
More biochemical and larval taxonomic research
would probably have obliterated some of the “sub-
species” and shown them to be species in their own
right. Here is certainly a field open for further
research.
The keys are in general good, short, useful and
working. The larval key to the last instar larvae of
Orgvia, however, leaves a question open: how can
there be a description ofa larva of a “subspecies” when
thee somscalled@ sty pels 1OlethisiessubSpeciesia mS) as
questionable as can be (“/eucostigma plagiata’) and
the larvae in question just do not yield adults like this
“type”? Otherwise this key is helpful and correct.
Certainly the most difficult genus is Dasychira.
LS)
204 THE CANADIAN FIELD-NATURALIST
Here Ferguson, who was especially involved for
decades with this genus, can proudly show the fruits of
long labor. The explications about the generic name
now make Dasychira finally definite for our species,
and we have to congratulate Ferguson for his success
in re-finding D. tephra. From this point of departure
everything else falls into place. We are also thankful
that he was able to confirm and give new meaning to
the names and taxa so familiar to all of us but so often
doubted by overzealous workers over many years. I
mean obliquata, cinnamomea, atrivenosa, meridion-
alis, basiflava, leucophaea, dorsipennata, plagiata,
grisefacta. Other names like aridensis, parallela,
lemmeri, pini disappeared after having been the
reason for much misunderstanding in the past.
The other difficult genus in the family is Orgyia.
Regrettably one cannot say what we just said about
Dasychira in this case. There are many needless
mistakes in Ferguson’s treatment. There are also good
points, and these are the following:
(a) the confirmation of the fact that all North
American species belong to one only well-defined
genus, Orgyia.
(6) the confirmation of O. cana as species in its own
right. This, however, is not a “revised status,” as it was
already treated as such in Entomologische Zeitschrift
83: 12, 1.e., in 1973, a publication which is left out of
the literature quoted. The problem with the associa-
tion of the larvae in this group would have been
completely and easily solved if Ferguson had used the
preserved and associated specimens of the McFarland
rearings in the Los Angeles County Museum of
Natural History. It has to be pointed out again that
MONA authors seem to rely too exclusively on the
collections of the United States National Museum, as
already Rindge objected in Journal of the Lepi-
dopteran Society 28: 4;
(c) giving taxonomic status to these puzzling very
large California Orgyia (magna). Further research 1s
here very much necessary:
(d@) the detection of the old Boisduval specimen which
Guérin-Méneville figured and used for the name of O.
detrita, in the collection of the United States National
Museum. It should be remarked that “Degens Bd.
Am. B.” is not “apparently... an unpublished name”
but is Latin for “coming from Boisduval America
Borealis” (the Latin verb is dego, -i, -ere);
(e) the additional knowledge about O. falcata and its
larva.
On the negative side there have to be mentioned the
following points:
(a) the laconic statement “the female genitalia have
not been studied.” Besides the fact that they have been
very extensively studied in several issues of Ento-
mogische Zeitschrift and are especially in the genus
Orgvia of decisive taxonomic importance, in a
Vol. 93
“definite” presentation of North American moths, this
is not quite understandable;
(b) the omission of all observations about pupae is
likewise not quite understandable, the more so as the
form of the pupa is also species characteristic;
(c) the omission of any study of the eggs by means of
the scanning electron microscope (SEM) and electro-
phoresis and also of allimportant larval characters, as
mandibles, ocelli, setae on thoracic legs, headcap-
sules, surface of exterior cuticle;
(d) the claiming of a “new synonymy” for O. definita
kendelli on p. 75 while this synonymy was long
established in Entomologische Zeitschrift 83: 14
(Definita kendelli = leucographa), 1.e., in 1973;
(e) the establishing of “new status” in the rindgei/
leuschneri complex while Chua et al. had already
established and published in Journal of Research on
the Lepidoptera 15: 4 (1976) specific status for O.
rindgei. This has been also repeated by Riotte in
Entomologische Zeitschrift 87: 3 (1977).
Some special words have to be said on the “enfant
terrible” of the whole fascicle: O. /euwcostigma. Looked
at as Ferguson presents it, it well makes sense. The
fact, however, is that the realities are quite other: (a)
The aedoeagus which is said “may be at once
distinguished from all other species in that it is
apically tapered to a point” is blunt and not tapered to
a point as SEM micrographs show. (6) The (not
studied) female genitalia would have helped to an
other and correct classification. (c) Use of the earlier
published results of egg electrophoresis would have
shown that indeed /eucostigma is sympatric to the
extreme with wardi which in no way 1s a “subspecies”
or “synonym” of anything. (d) Larval structures, if
they had been used, would have shown the same (they
also were previously published). (e) To use the so
called “Walker type” of Acypha plagiata, abdomen-
less as it is and without any locality label, as type fora
“subspecies” of /ewcostigma in Nova Scotia, replacing
wardi, 1s at best absurd. Insect pins may be convincing
sometimes; in a case like this, certainly not. The
Walker specimen could be used for the real /euco-
stigma in Nova Scotia but I can see no need for this.
Competent workers in the British Museum came toa
quite other evaluation of the Walker specimen and
placed it together with /eucostigma from Wisconsin as
best match. Therefore, one should list the Walker type
as “incertae sedis.” One good thing at least should be
mentioned, however: with Abbot’s larval painting of
leucostigma declared as lectotype of the species, we
have now a good basis for it. We also think that the
male figure of Abbot’s /ewcostigma is really definita
(many of Abbot’s plates are mixed with non-
conspecific creatures). Very good also is Ferguson’s
final clearing up of the /ewcographa Geyer problem:
we have nothing against his treatment here.
1979
The plates in this volume are of the expected quality
and with the richness of depicted specimens very
useful. The only thing we miss Is a single specimen of
Orgvia wardi from the type locality, Prospect Road,
N.S., perhaps best together with a /eucostigma from
the same locality to show the difference. The Nova
Scotia specimens figured are all /eucostigma.
In the literature quoted we miss the important
paper by Chua et al., 1976, Investigation of selected
species of the genus Orgyvia (Lymantriidae) using
isoelectrofocusing in thin layer polyacrylamide gel
(Journal of Research on the Lepidoptera 15(4): 215—
224) as well as Riotte, 1973, Uber Orgvia (O.) gulosa
und Orgyvia (O.) cana (Lep.: Lymantriidae) (Ento-
mologische Zeitschrift 83(12): 129-140) and by the
Hawks, Falcons and Falconry
By Frank L. Beebe. 1976. Hancock House, Saanichton,
British Columbia. 320 pp., illus. $12.95.
Any author attempting to write an original book on
birds of prey or falconry today, faces a difficult task,
but Beebe certainly does offer something new in this
most recent book. Perhaps the book should be
retitled “Hawks, Falcons, Falconry and Human
Attitudes”; through the entire book, there pervades
an undertone of bitterness towards an anti-falconry
movement, which culminates in the final chapter,
“The endangering of the Peregrine: A study in
environmental strategy.” A reader cannot help but be
impressed by the collection of facts which, according
to the author, led to classifying the Peregrine Falcon
as an endangered species. After seven years of intense
involvement with birds of prey and their human
admirers, be they scientists, bird-watchers, falconers,
photographers, or others, I find it difficult to believe
that these facts constitute a case for the premeditated
abolition of falconry.
Biologists will be disappointed by the rather brief
treatment of general raptor biology but Beebe
demonstrates a good grasp of raptor behavior. His
observations and theory on the adaptive significance
of a reversible hallux, prone sleeping position, and
very deep sleep associated with Arctic-nesting Gyr-
falcons (p. 167) is interesting and certainly warrants
further investigation. On p. 180, the author provides
some fascinating insight into the influence of geo-
graphical landforms on hunting strategies of river-
nesting Peregrine Falcons.
The descriptions for each species covered in the
book are fairly standard, but tend to lean toward the
western areas of Canada with which the author is
clearly most familiar. Much, if not all, of this material
has been reproduced from his earlier published work
BOOK REVIEWS
205
same author, 1977, Abschliessende Bemerkungen zu
den Studien uber nordamerikanische Arten der
Gattung Orgvia (Lepidoptera: Lymantriidae) (Ento-
mologische Zeitschrift 87(3): 9-12, concerning defi-
nita, rindgei, and leucographa = detrita).
The criticisms, however, do not interfere with the
great value of the fascicle otherwise. We certainly
welcome its final appearance and wish it good and
longlasting success.
URC2 EMR TOnmeE
Department of Entomology, B.P. Bishop Museum, P.O.
Box 6037, Honolulu, Hawai, USA 96818 and Royal
Ontario Museum, Toronto, Ontario
“Field studies of the Falconiformes of British
Columbia,” No. 17 Occasional Paper Series, British
Columbia Provincial Museum. The author is some-
what inconsistent in his descriptions of the various
subspecies, providing them for some species, but not
for others.
The section on falconry techniques should prove
extremely useful to falconers, as Beebe has chosen to
present detailed accounts of certain aspects of this art
rather than give a shallow description of falconry as a
whole. For example, he has given a thorough
description of how to fly a bird to the lure (p. 239).
Having conducted research on kestrel reproduction,
I must dispute the author’s statements regarding the
timing of the breeding for this species (p. 132). Beebe
claims that “kestrels arrive on their breeding terri-
tories in April” and that “eggs are produced,
depending on latitude, from late May to mid-June.”
In southern Quebec at least, kestrels generally arrive
as early as the latter half of March and lay eggs during
the last two weeks of April and the first two weeks of
May, somewhat dependent on weather conditions
from year to year. Furthermore, my studies and those
of others (Bent, A. C. 1938. Life histories of North
American birds of prey. Part 2: Orders Falconiformes
and Strigiformes. Dover Publications, New York. 482
pp.; Balgooyen, T. G. 1976. Behavior and ecology of
the American Kestrel (Falco sparverius) in the Sierra
Nevada of California. University of California Pub-
lications in Zoology 103: 1-83), give an average egg
size of kestrels as approximately 35 X 28 mm rather
than the 39 X 29 mm Beebe reported.
The first-year mortality rate of 90% quoted in this
book for birds of prey on p. 207 is surely the highest
ever reported in the literature. Also, this reviewer was
206
not aware that the seal hunt had been closed (see p.
298).
There is much unnecessary repetition in photo-
graphs and an overabundance of head studies, some
of very poor quality, e.g., Peale’s Falcon, p. 170. The
ink drawings, with some exceptions, are quite well
done, as well as functional. The flight silhouettes
throughout the book are excellent. One illustration,
that of a falconer’s knot, is apparently missing on p.
193, and on p. 239 the labels on the two diagrams
apparently have been assigned incorrectly.
These latter two errors exemplify the poor editing
found throughout the book. More than 50 typo-
graphical and spelling errors were located in the text
and there was inconsistent use of names, i.e., kestrel
versus sparrowhawk, and even spelling, 1.e., grey
versus gray. References were not presented in a
consistent manner and sometimes are incorrect, e.g.,
Morlan W. Nelson (1965), p. 152. Much space is
Fishes of the World
By J.S. Nelson. 1976. Wiley, New York. xii+416 pp.,
illus. US $26.00.
The author wrote this book “to present a modern
introductory systematic treatment of all major fish
groups.” It is intended to serve as a reference for
professionals and as a text for fish systematics
courses. This book, therefore, was doomed to failure
before it was written, and ensured of success before it
was off the press. Fish systematics is ina state of rapid
change and the healthy diversity of opinion on various
aspects of fish systematics assures that some of the
included material is outdated, but the need for any
recent synthesis is obvious to anyone trying to teach
the subject.
Following the prefatory remarks, the text includes
an Introduction which comments on numbers of
fishes, their diversity, distribution, and classificatory
schemes. Then follows a classification of the phylum
Chordata in which fishes are treated to at least the
familial level. Appendix I is a checklist of extant
classes, orders, suborders, and families. Appendix 2 is
a series of 45 distribution maps of families of fishes.
There is an extensive bibliography and the Index
includes the genera mentioned in the text.
Nelson considers six classes of fishes (four extant)
comprising 46 orders and 450 families. An outline
drawing of a “typical” representative of each of most
families is included. Characteristics for each included
family or higher taxon are usually included, although
they are not always complete. Recent revisionary
papers on the orders are mentioned and comparisons
THE CANADIAN FIELD-NATURALIST
Vol. 93
occupied by overly emphatic titles in the first nine
sections where only brief treatment of the subjects
follows.
There is no doubt that Beebe has had much
experience with both wild and captive birds of
prey and for this reason, I am more inclined to
recommend his other book, “Field studies of the
Falconiformes of British Columbia” for raptor en-
thusiasts not especially interested in falconry. Fal-
coners from coast to coast, however, will likely
applaud Beebe’s efforts in this book to speak on their
behalf and it is to this audience that I heartily
recommend this book.
DAVID M. BIRD
Raptor Research, Department of Renewable Resources,
Macdonald Campus of McGill University, Ste. Anne de
Bellevue, Quebec H9X 3MI
with previous classificatory syntheses are included.
The number of genera and approximate number of
species in each order and family are given, and insome
cases these taxa are mentioned.
There are, of course, the inevitable minor errors:
e.g., there is only one species of Zanclus, not two;
Terapon is misspelled Therapon: and a few of more
consequence, e.g., the retention of the families
Rosauridae and Kasidorodae. For a volume of its size
and complexity of nomenclature, there are re-
markably few typographical errors and the references
to literature through 1973 are quite complete. In-
evitably this reference will be compared to the recent
book by Lindberg which, unfortunately, in the
English translation has the same title. The two books
complement one another: Nelson’s classification is
more modern, has some distribution maps, and costs
less than Lindberg’s, which has better illustrations, a
more extensive list of references and, of course, keys
to families.
By and large Nelson has achieved what he set out to
do, and despite differences of opinion one may have
with the classification he uses or the philosophy by
which he arrived at it, this is a most useful text. I
recommend it highly to all serious students of fishes.
C. GRUCHY
National Museum of Natural Sciences, Ottawa, Ontario
KIA OM8
1979
BOOK REVIEWS
207
Nesting Ecology of Canada Geese in the Hudson Bay Lowlands of Ontario:
evolution and population regulation
By Dennis G. Raveling and Harry G. Lumsden. 1977.
Ontario Fish and Wildlife Research Report Number
98. Ontario Government Bookstore, Toronto. v +
77 pp. $3.50.
The report is divided into two major sections,
nesting ecology and population limitations. Objec-
tives for the study of the nesting ecology of Canada
Geese of the Mississippi Valley Population were to
examine apparent optimum and accomplished repro-
ductive rates, habitat preferences, and density and
spacing of nests in relation to the regulation of
population size. The primary objective for the second
portion of this report was to examine nesting ecology
to provide insight into the factors limiting goose
numbers. These objectives have certainly been ful-
filled. The study has direct management implications
and extends our understanding of the evolution of
waterfowl adaptations to their environment.
The 159-square-mile study area located west of
James Bay was examined intensively, by walking and
helicopter searches, in 1967-1969 for nesting geese.
The researchers were fortunate in having a “late” year,
an “early” year, and an “average” year, making for
some interesting year-to-year comparisons. Among
the many factors measured that relate to nesting
ecology were habitat preferences, nest site location,
nest phenology, nest density and spacing, clutch size,
nesting success, predator influences, fertility, and
goose behavior. An especially interesting facet of the
study involved the development of a model of body
weight dynamics of adult females in spring that
calculated the caloric cost of migration, maintenance,
Birds in Boreal Canada
By Anthony J. Erskine. 1977. Canadian Wildlife Service,
Report Series 41. Supply and Services Canada,
Ottawa. 71 pp., illus. $5 in Canada; $6 elsewhere.
This book presents “an overall review of the boreal
avifauna, its composition, evolution, and prospects
for survival.” For eight summers, Erskine made
quantitative studies of boreal birds, from New
Brunswick to British Columbia, utilizing 40-hectare
(16-acre) plots, each of which he visited three or more
times.
Merriam’s concept of a boreal life zone is seen to be
a misleading oversimplification. Erskine shows how
bird species composition varies greatly between black
spruce, tamarack, balsam fir, and aspen. Further
subcategories such as ‘shrubs after burns,’ ‘farms,’ and
‘balsam poplar of flood-plain forest,’ allow even better
understanding of bird distribution and abundance.
egg laying, and incubation and predicted weights and
control of clutch size. Energy stores accumulated
during spring migration were shown to be essential to
flight to nesting range, waiting for nest sites to become
snow free, egg formation, and maintenance through
incubation. Clutches were largest in an “early” year
and in early nests within a year. The mechanism of
cessation of egg laying appears to be the exhaustion of
accumulated reserves to a “winter” weight level.
Winter mortality was clearly the major limiting factor
for Mississippi Valley Population geese, and this
mortality was overwhelmingly accounted for by
hunting.
The report contains 54 tables and 34 figures, all of
which are neat and clear. Several black-and-white
aerial photographs illustrate the various habitat types
in the study area. The text is lucid and very readable.
In fact, the only negative aspect of the entire report is
its slightly awkward size, being 8 X 11".”' rather than
8", X 11”. Anyone interested in avian nesting ecology
and in the importance of well-designed and thorough
field studies will find this report to be an excellent
source.
NOEL J. CUTRIGHT
Wisconsin Electric Power Co., 231 W. Michigan, Milwaukee
WI 53201
'Book Review Editors Note: 20 cm X 30cm as result of
Ontario Government Metrification program.
Although the American Robin and the White-
throated Sparrow occupy no less than ten of these
habitat niches, the Black-and-White Warbler is listed
for only one (poplar-birch forest). The Tennessee
Warbler is listed for broad-leaved understory of
spruce, for jack pine, for alder understory of tamarack
bog forest, for young aspen, and openings (western),
but is rare in pure spruce. In stands where balsam fir
predominates, one may expect to hear the Winter
Wren, Blackburnian and Black-throated Green War-
blers, with Bay-breasted Warblers in the more mature
stands, and Magnolia Warblers in the younger
growth, while the Ruffed Grouse and Blue Jay replace
the Spruce Grouse and Gray Jay. One should look for
Nashville Warblers in tamarack bog forest.
Inevitably others will disagree with Erskine on
some minor points. I would not agree that the Red-
208
tailed Hawk is more characteristic of conifer than of
broad-leaved forests. I would add Sharp-tailed
Grouse, Black-billed Magpie, and House Sparrow to
the list of birds characteristic of northern farms, the
Great Crested Flycatcher, Cedar Waxwing, and
Connecticut Warbler to the list for aspen forest,
especially in parkland areas, and the Vesper Sparrow
to the list for more southerly jack pine forests. I do not
agree that Least Sandpipers breed in boreal fens at
this latitude.
Erskine contends that birds of the boreal forest
have attracted less study than those of any other part
of Canada. Nevertheless, his is a personal project that
fails to review many published studies from the
THE CANADIAN FIELD-NATURALIST
Vol. 93
region; Flack’s review of aspen parkland birds is
mentioned but some of Flack’s important conclusions
are ignored.
This is a landmark publication. It heralds a new
recognition by the Canadian Wildlife Service of its
responsibility for “dickey birds” and their ecology. It
contains a wealth of ecologic and distributional data,
brilliantly analyzed and synthesized. It demands
careful and repeated perusal by everyone interested in
boreal birds.
C. STUART HOUSTON
863 University Drive, Saskatoon, Saskatchewan S7N 0J8
Lexique anglais-francais, Termes techniques 4 I Usage des Biologistes
By Jean Vaillancourt. 1978. Editions de ’Université d’Ot-
tawa, Ottawa. 427 pp. Paperback $12.00.
Biologists, like the animals they study, must look
beyond linguistic and national boundaries, to be
aware of developments in their field. This lexicon, of a
high degree of completeness, will help the biologist
reap the full benefits of discoveries in two of the
world’s most important languages.
The lexicon consists of two principal parts. The first
and largest is an English-French lexicon with the
entries listed alphabetically in English and numbered
consecutively. The emphasis is thus on providing the
equivalent word in the other language rather than a
definition. The second section is an alphabetical index
of French terms keyed by number to the first section.
This approach permitted condensation of the text and
reduction in cost without loss in utility.
The English-French lexicon is laid out in the
following manner: the English word in boldface type,
which permits the word to be easily picked out,
followed by one or more classificatory abbreviations
such as Anat., Biol., Zool., Océanogr., Méd., Bot.,
Ichtyol., Ornith., Micr., etc.; the French synonym or
synonyms; the plural form (only when irregular); and
the gender.
The only other even roughly comparable sources
the reviewer is aware of are the following: Glossaire de
biologie animale by Roger Husson (1970, Gauthier-
Villars, Paris), which is unilingual and gives about
2500 definitions; the Dictionnaire frangais-anglais,
anglais-frangais des termes médicaux et biologiques
by Pierre Lépine and Philip R. Peacock (1974,
Flammarion, Paris), having some 6500 words; and the
Dictionary of biology by Gunther Haensch and
Giselin Haberkamp de Anton (1976, Elsevier Sci-
entific Publishing Company, Amsterdam), with syn-
onyms in four languages for about 9800 terms. (There
are also a number of unilingual English biological
dictionaries.) None of these give comparable cover-
age. To check the thoroughness of coverage the first
25 biological terms found in the preceding diction-
aries were checked to see whether they were included
in Vaillancourt’s lexicon. Of the 25 terms in all three,
about 90% were found in Vaillancourt. The reviewer
has a rather specialized manuscript dictionary of
ichthyology. Even in this case about half these
specialized terms were found in Vaillancourt. The
coverage 1s clearly very good. Over 10 400 terms are
included.
The disciplines covered include bacteriology, bot-
any, zoology, marine biology, anatomy, systematics,
evolution, ecology, limnology, oceanography, genet-
ics, paleontology, pharmacology, and physiology as
well as related terms in geology and geography.
In a discipline as broad as biology it would be
impractical to include every term. Only a few
omissions were noted — neotype (néotype), meristic
(méristique), and ray in the sense of fin ray (rayon),
although ray in the sense of a kind of fish was
included. Ichthyologist and fish(eries) biologist are
equated, although in general use the first is usually
restricted to the theoretical student of systematics,
anatomy, evolution, ecology, and zoogeography of
fishes, as opposed to the applied management of
fishes; however, these are minor points.
I wholeheartedly recommend the addition of this
book to your biological library. There have been
many times that I wished I had a comparable
authoritative source book. Vaillancourt is to be
congratulated for his breadth of coverage which will
make it invaluable for the student and professional.
Don E. MCALLISTER
National Museum of Natural Sciences, National Museums
of Canada, Ottawa, Ontario KIA 0M8
IO79
BOOK REVIEWS
209
An Introduction to the Aquatic Insects of North America
Edited by R.W. Merritt and K.W. Cummins. 1978.
Kendal/ Hunt, Dubuque, Iowa. 441 pp., illus. US $18.95.
Nearly twenty years have passed since a new
comprehensive reference work on aquatic insects has
been published. Although the publication of this book
dealing with aquatic entomology will not render any
of the existing texts obsolete, it is a long awaited up-
to-date synthesis of the taxonomic status of the
various groups of aquatic insects. The greatest
improvement over existing books of this kind is the
shift in emphasis from the traditionally taxonomic
theme to one that includes extensive ecological
information.
The taxonomic treatment of the insect orders
presented in the text has been restricted for the most
part to the familial level. Only a few of the smaller
orders such as the Megaloptera and the Neuroptera,
and a few families of aquatic Diptera, notably the
Tipulidae,Culicidae, and Simuliidae, are dealt with at
the generic level in the keys. The opinion of the editors
was that a single reference work treating all major life
stages to the generic level, as well as including the
ecological data, would be of prohibitive size. Pre-
sumably the inclusion of the extensive ecological
information for the orders has been done at the
expense of the taxonomic sections to provide a more
encompassing review. This decision by the editors is
commendable for it has resulted in a wide range of
information available in a single text. In view of the
frequent changes that occur at the generic and specific
levels, the restriction of the taxonomic information to
the family level by the editors should also result in a
book less subject to the need for major revisions. In
addition each section contains a listing of the major
references to more detailed taxonomic studies. Thus
the lack of generic keys has little impact on the
usefulness of the book.
Although the taxonomic sections generally deal
with the organisms at the family level, most of the
genera are mentioned in the ecological treatment of
each order. The ecological data has been compiled
into tables and the editors have maintained a
consistency throughout the book while including as
much ecological information as possible in a readily
obtainable form. This is especially significant. con-
sidering the diversity of contributors involved in the
preparation of the text. The reader need only find the
genus of interest and read across the page to determine
its habitat, behavioral characteristics, trophic rela-
tionships, and distribution in North America. As well,
references to specific ecological studies are also
included if more information is required.
Additional topics pertaining to aquatic entomology
serve as introductory to the systematic and ecological
sections on the insect orders. The section dealing with
sampling methods and equipment deserves special
mention. The full range of aquatic habitat types is
presented in tabular form with a listing of the
sampling methods and equipment considered most
suitable for each habitat type. The references to
pertinent literature that are contained in this section
appear to have been carefully selected to include both
proponents and critics of the various methods
outlined in the tables. The most commonly used
equipment is well illustrated by diagrams, and a list of
suppliers, complete with addresses is provided in the
text. Canadian readers, however, will be disappointed
that this list contains only suppliers in the United
States and does not mention Canadian distributors.
The chapter dealing with the general morphology of
the aquatic insects is, like the rest of the book, well
illustrated by diagrams. Unfortunately, only the
stoneflies are illustrated in the immature stage, which
in a book geared towards students, provides little
guidance for those attempting to work with highly
specialized forms suchas the larvae of aquatic diptera.
A generalized treatment of phylogenetic relation-
ships and evolutionary adaptations of aquatic insects
completes the non-systematic sections of the book.
The book appears to be aimed towards the amateur
collector and the beginning student of aquatic
entomology, but its use is by no means restricted to
this group. Although the generalized treatment of the
topics contained in the text clearly demonstrates this
bias, the work is also an excellent source book that the
professional can draw from. The editors have defi-
nitely succeeded in producing a work that will be of
use to all levels of interest and expertise in aquatic
entomology for many years.
REIN JAAGUMAGI
Department of Entomology and Invertebrate Zoology,
Royal Ontario Museum, Toronto, Ontario MS5S 2C6
210
Nichoirs d’ oiseaux
Par Raymond Cayouette, illustrations de Jean-Luc
Grondin. 1978. La Société zoologique de Québec,
Charlesbourg, Québec. 36 pp. $4.
Cette plaquette brochée de 36 pages a des
dimensions qui sortent de l’ordinaire (25,5 x 20,0 cm),
mais qui contribuent a mettre en relief la valeur
générale de Tlouvrage. La composition et la
présentation graphiques sont trés agréables et de
bon gout. Le papier est de bonne qualité et
limpression est soignée. La reproduction des planches
en couleurs compte parmi les meilleures que j’aie eu
Yoccasion d’examiner au cours des derni€res années
et rend justice au talent de l’artiste.
L’ouvrage comprend cinq parties principales qui
varient en importance.
L’introduction et les généralités occupent les huit
premieres pages et on trouve en neuviéme page une
liste des espéces qui utilisent des nichoirs. Les auteurs
traitent en détail de vingt espéces d’oiseaux nicheurs
dans les vingt pages suivantes, et des modéles de
nichoirs accompagnés de dimensions apparaissent a
la fin de ’ouvrage.
Ce qui frappe surtout dans cette plaquette c’est la
finesse des illustrations, c’est du Jean-Luc
Grondin a son meilleur. Grace a ce travail Grondin
s’affirme parmi les meilleurs artistes-naturalistes du
pays. Son style est sir et il maitrise trés bien sa
technique. I] concilie a la fois le souci du détail et un
grand sens artistique et démontre qu'il est aussia l’'aise
avec les illustrations en couleurs qu’avec le dessin au
trait qu'il a si heureusement utilisé antérieurement.
Le texte Raymond Cayouette est clair, précis et bien
rédigé. On y trouve une quantité de détails
intéressants et généralement précis sur la nidification
des oiseaux en général et sur les 20 espéces qui
sont traitées dans le texte. Ces derniéres sont
accompagnées dillustrations. En page 5, l’auteur
mentionne que l’Hirondelle pourprée “a diminué de
Birds in Peril: a guide to the endangered birds
By John P. S. Mackenzie. 1977. Pagurian Press, Toronto.
191 pp., illus. Hardback $14.95, paper $7.95.
I have often felt that an author who attempted to
write a topical book on an environmental issue did so
at considerable risk, for new information Is constantly
being revealed that dates the author’s effort or
contradicts his work. Endangered species has
especially been the topic of considerable verbiage,
much of it ill-informed. Despite these obstacles, John
Mackenzie has made a worthwhile effort in bringing
to the public’s attention the plight of some endangered
or threatened birds.
THE CANADIAN FIELD-NATURALIST
Vol. 93
facon évidente la quantité de moustiques” dans une
région des Etats-Unis. Or il s’est avéré, a la suite
d’études récentes, que lHirondelle pourprée con-
somme peu de moustiques! On a quelquefois con-
fondu “couvée” avec “ponte.” Ponte désigne toujours
l'ensemble des oeufs pondus par un oliseau.
J’ai constamment noté que les longueurs des
oiseaux different appréciablement, dans certains cas,
des longueurs reconnues pour ces espeéces. Les
périodes de couvaison données sous le vocable “Durée
de l’incubation” ne sont pas toujours en accord avec
celles reconnues dans des études de ces espéces. De
méme le réle des adultes dans la couvaison, tel
qu’indiqué dans le texte, n’est pas toujours conforme
aux données fournies ailleurs! Les aires de nidification
au Québec sont générales, mais précises sauf dans le
cas de ’Etourneau sansonnet que lon fait nicher a
Fort-Chimo!
Les dimensions recommandées des_ nichoirs
apparaissent dans un tableau ingénieux en page 33 et
dérogent peu des dimensions recommandées par
plusieurs auteurs. Les modeéles de nichoirs suggérés en
pages 34, 35 et 36 ne sont pas innovateurs, mais
suffisent pour qui veut se donner la peine de les
construire pour le plus grand bénéfice des oiseaux qui
les utiliseront.
En résumé, cette petite brochure, malgré quelques
imprécisions de texte, est un excellent achat, méme a
$4.00, a cause de la beauté des illustrations et des
renseignements qu’elle renferme. C’est sans réserve
que j’en recommande lachat a tous ceux qui
s’intéressent aux Oiseaux et qui veulent leur offrir des
sites de nidification supplémentaires.
HENRI OQUELLET
Musée national des Sciences naturelles, Musées nationaux
du Canada, Ottawa, Ontario. KIA 0M8
of Canada and the United States
The book relates the history of 20 species or sub-
species of birds in North America, how they became
threatened, and what steps are being taken to help
prevent their extinction. Refreshingly, stories have
been included to illustrate that intelligent wildlife
research and management can help bring a species
back to a reasonable population.
Terry Shortt produced twenty color paintings for
the book. They have been delicately reproduced and
capture the spirit of his art. Additional line drawings
are throughout. As usual, the work of Terry Shortt is
superb. Having just visited an exhibit of his work that
1979
included the original color paintings for the book, I
recommend the book for Shortt’s paintings alone.
The introductory pages are clearly but simply
presented with no new insights or information
offered. A discussion on the complex factors that
usually work together to cause a species to become
threatened would have been helpful. For example,
should threatened wildlife be managed on a species,
sub-species, or population level? Or, should an animal
be considered endangered when it is merely occurring
on the fringe of its range? The initial discussion was
inadequate.
Each discussion is broken into sub-headings:
history, decline, life cycle, aspects of biology and
behavior, what is being done and the future. This is a
good feature, for it provides a wide range of
information that can easily be found for each species.
Much more could have been added and the reader
should not view the text as definitive accounts.
Occasionally the writing is unclear. On page 12, for
example, the comment, “commercial interests such as
mining, oil, gas . . . support environmental studies”
“financed a multitude of environmental studies on all
aspects.” True, corporations have undertaken many
studies but we cannot lightly dismiss corporate
BOTANY
Manual of the Vascular Plants of Wyoming
By Robert D. Dorn. 1977. Garland Publishing Inc., New
York and London. 1498 pp. (2 volumes). US $95.00.
These two volumes, if they had been typeset rather
than typewritten double-spaced, and often with large
blank spaces on the pages, could have been condensed
into one easily handled volume.
The author has deliberately not included infra-
specific taxa, some annual grainfield weeds, species
growing only under irrigation conditions suchas lawn
weeds, and species to be expected but not yet found in
the state. One cannot argue about a decision to
produce a conservative treatment, and as well not to
include extraneous taxa. It does seem unfortunate,
however, that a large part of the introduced flora has
been ignored. This is particularly so, because students
who will test the keys will probably do so on plants
gathered from about townsites and college campuses,
and thus be frustrated because the correct answers are
not always forthcoming.
The order of the flora is strictly alphabetical within
the three large groupings of Pteridophytes, Gymno-
sperms, and Angiosperms. Descriptions are relatively
short: habitat information is adequate; distributions
BOOK REVIEWS PAU
responsibility for on page 22, the author mentions the
hazards of oil tanker traffic, which will be greatly
increased when the pipeline is complete. As far as lam
concerned, the jury is still out on both government
and corporate responsibility towards the environ-
ment. Wildlife is just not perceived by decision-
makers as a competing resource.
The inclusion of success stories in the management
of endangered species is a welcome addition. Dis-
cussions on the Trumpeter Swan, Hudson Godwit
and others is helpful. I found too much emphasis
placed on the work being done by the United States
Fish and Wildlife Service, although this may reflect
the most ambitious program. I had the feeling at times
that the “what is being done” section was overkilled
and tended to be repetitive.
One can quarrel about many “nits” especially by a
reviewer who works in the wildlife management field,
but on balance, the book is well put together and
provides a good source of information for the
layreader.
GERALD B. MCKEATING
14 DeSavery Crescent, Toronto, Ontario M4S 2L2
are given only for the state of Wyoming and
sometimes only by the name of a county; synonomy is
minimal; the few line drawings scattered throughout
are by Jane L. Dorn.
An appendix near the end of the second volume
includes a page on geography and climate, a map of
the state depicting counties, sections on historical
vegetation, present vegetation, natural vegetation
types, selected references, rare and endangered spe-
cies, early collectors in Wyoming, and a summary
giving a breakdown by family of the 605 genera and
2144 species treated. This might perhaps better have
been placed with the introductory material in the first
volume.
The high cost, coupled with the bulkiness of this
work, will undoubtedly mean that few students, for
whom it was written, will put it on their bookshelves.
WILLIAM J. CODY
Research Institute, Agriculture Canada.
KIA 0C6
Biosystematics
Ottawa, Ontario
D2 THE CANADIAN FIELD-NATURALIST
Vol. 93
A Provisional Checklist of Species for Flora North America (Revised)
Edited by S.G. Shetler and L. E. Skog. 1978. Missouri
Botanical Garden Monographs in Systematic Botany
Volume |: 1-199. Paper US $6.50.
This revised checklist which includes 16 274 species,
2350 genera and 233 families is a tangible result of the
efforts of many individuals, and particularly of
Shetler, towards what was eventually to have pro-
duced a Flora of North America. It 1s indeed a step
towards such a flora, and is a document from which a
general review and revision of the flora can begin.
The area covered is that part of North America
north of the Mexican border, including Greenland.
Included are names of taxa, authorities, coded
information of four categories of plant character-
istics, range given by 16 regions, and the source of the
information. There is no synonymy. The computer
printout has been photoreduced in order to incor-
porate more information on each page, and this has
made legibility somewhat more difficult.
WILLIAM J. CODY
Biosystematics Research Institute, Agriculture Canada,
Ottawa, Ontario KIA 0C6
Garden Spice and Wild Pot-herbs. An American Herbal
By W.C. Muenscher and M.A. Rice. 1978. Cornell Uni-
versity Press, Ithaca, New York. 213 pp., illus.
US $16.95.
This book contains accounts, botanical, culinary,
and horticultural, of approximately 70 plants, pri-
marily cultivated plants, used as herbs and spices. An
illustration of each plant is provided, showing the
stage at which the particular plant should be
harvested. The illustrations themselves have been
prepared from the original wood-cuts by Elfriede
Abbe. Included in the botanical aspects of the book
are short accounts of the plant families represented as
well as brief descriptions of each of the species
included. Scientific names are given, with some
frequently used synonyms being listed in the appen-
dix, as well as common English, French, and Italian
names. The culinary notes are rather general but
include information on the usual uses of various
portions of the plants. I suspect that cooking has a
“common-name problem” similar to that which led to
the establishment of scientific nomenclature: horse-
radish sauce is described as “an alcoholic infusion” to
my great surprise. I can only suspect that Muenscher
and I know different sauces by the same name.
Medicinal and nutritional notes are also included,
the former I suspect being primarily of the folk-
medicine genre. It is unfortunate that the glossary is
restricted to botanical terms, for many of those used in
the medicinal portion are probably unfamiliar to the
majority of readers. What does a carminative drink
do?
For those with gardens, but particularly those with
gardens in southern Canada, the horticultural in-
formation will be of interest. In this regard, the
hardiness table, showing survival rates at Ithaca, New
York, for the winter of 1942-43, offers more concrete
information than is usually available.
My own reaction to the type-face used, Goudy
Kennedy Bold and Italic, was negative: it seemed hard
on the eyes. On the other hand I think it did help to
induce a relaxed, browsing mood and this is, | would
say, a book for browsers rather than readers. It
contains a lot of fascinating information and I
anticipate dipping into it fairly frequently both when |
am planning a meal and when asked a question by a
home gardener interested in growing herbs and spices.
It is a reprint of the original 1955 edition.
MARY E. BARKWORTH
Biosystematics Research Institute, Agriculture Canada,
Ottawa, Ontario KIA 0C6
How to Identify Grasses and Grasslike Plants (Sedges and Rushes)
By H. D. Harrington. 1977. Swallow Press Inc., Chicago,
Illinois. 142 pp., illus. US $3.95.
Grasses, sedges, and rushes are often not even
considered “proper” flowering plants by amateurs and
some beginning botanists. This erroneous impression
is reinforced by non-technical identification manuals
which totally ignore the three families. The absence of
an attractive corolla can apparently only be accepted
in trees, which, of course, do not have ffowers as in
“How to identify the flowers of ABC.”
Undoubtedly, learning to identify and eventually to
recognize the grasses, sedges, and rushes is not
179
straightforward. The structures involved are some-
what different and generally rather small. Neverthe-
less so much of our vascular plant flora belongs to
these three groups that the botanist, whether amateur
or professional, who ignores them or lumps them into
a few genera ignores a substantial proportion of the
plants in the area.
Harrington’s book is not an identification manual.
It is, however, an illustrated dictionary and advice
manual that anyone tackling the three families for the
first time will find an invaluable reference. Terms are
not only explained in words but are also illustrated
with line drawings. Moreover beginners will be parti-
cularly pleased to find that words such as ‘anther’ are
illustrated as well as ‘aphyllopodic.’
Most of the book is concerned with grasses (six
Flora of Alberta — a checklist
By D. F. Brunton. 1977. Alberta Department of Recrea-
tion, Parks and Wildlife, Edmonton. 43 pp. Free.
This small booklet represents a checklist of the
vascular plants of Alberta that has been compiled by
the author from E.H. Moss’s (1959) Flora of Alberta,
J. C. Packer and M. G. Dumais’ (1972) “Additions to
the Flora of Alberta” (Canadian Field-Naturalist 86:
269-274), and J. C. Packer’s (1974) A Supplement to
E. H. Moss’s Flora of Alberta. The listed species’
names and very abbreviated annotations concerning
their generalized distributions in Alberta (e.g., SW &
N) or rarity status (e.g., R) have been abstracted
entirely from these earlier sources. Thus, this booklet
does not represent either a nomenclatorial updating
or a reconsideration of the taxa composing the
Alberta flora. For most purposes, readers would seem
better advised to use the original sources, plus B.
Boivin’s (1968-72) Flora of the Prairie Provinces. For
more updated information on the status of Alberta’s
rare plants, readers should see G. W. Argus and D. J.
White’s (1977) “The Rare Vascular Plants of Alberta”
(Syllogeus, Number 17).
Despite their obvious limitations, I believe such
provincial, state or regional checklists of the flora may
serve useful purposes. Although not true in this case,
Freshwater Wetlands: ecological processes and
Edited by Ralph E. Good, Dennis F. Wigham, Robert L.
Simpson, and Crawford G. Jackson, Jr. 1978. Aca-
demic Press, New York. 378 pp., illus.
This book developed from a conference entitled
“Freshwater Marshes: Present Status, Future Needs.”
BOOK REVIEWS
713)
chapters) with one chapter each for sedges and rushes.
In addition, there is a chapter on collecting and
pressing and another listing identification manuals
that were available at the time of writing. Harrington
is consistently encouraging, straightforward, and
realistic. He states quite clearly that identification of
the three families is not easy but not impossible for the
non-professional taxonomist. In reading the book it is
quite evident that he has helped numerous students in
the past. He will continue to do so.
MARY E. BARKWORTH
Biosystematics Research Institute, Agriculture Canada,
Ottawa, Ontario KIA 0C6
checklists often provide the only coverage of the total
known flora of a region and represent an important
preliminary stage towards the ultimate production of
a floral manual with keys and descriptions. A
checklist provides a brief, easily consulted outline of
the flora and can be conveniently used to check off the
plants encountered in more local species’ inventories.
Space for just such a purpose has been provided in the
present booklet. It may prove to be most useful for
environmental consultants who are involved with
baseline inventory studies of particular areas for
impact assessments preceding planned developments.
Personally, I find such a checklist to be a convenient
source for quickly checking the spellings and author-
ships of species’ scientific names. I believe that the
practicality of the present checklist of the flora of
Alberta could have been improved by the inclusion of
at least the more important synonyms, especially -
those needed to correlate the species’ names used in
the floral manuals of Moss, Boivin, and others.
V. L. HARMS
Fraser Herbarium, University of Saskatchewan, Saskatoon
STN 0WO
management potential
The objective of both the book and the conference was
to consider all ecological aspects of wetland eco-
systems in addition to the production ecology of
marshes. This objective, of course, entailed an
expansion of the object of study from marshes to the
214
total spectrum of “wetlands.” Wetlands are generally
accepted to be the broadest classification category
comprising all lands inundated by standing water fora
sufficient portion of growing seasons to establish a
hydrophytic vegetation, or lands with a high water
table and an accumulation (generally accepted to be
greater than 30cm) of raw organic matter which
essentially provides the rooting medium for vege-
tation.
This development from conference to book eluci-
dates the genesis of the book’s title Freshwater
Wetlands but it in no way facilitates this reviewer’s
acceptance of the term. In the study of marshes,
freshwater and saline marshes are convenient and
practical subdivisions of the field of study. In the
study of wetlands, however, the acceptance of a
category of freshwater wetlands involves a flagrant
discrimination against a small group of saline wet-
lands. If generally adopted, this concept could inhibit
the investigation of the practical criteria which may be
used to distinguish the freshwater and saline sub-
divisions since workers would be reluctant to ap-
proach the ‘no man’s land’ between these two,
assumed, compartments.
An erroneous conclusion has been drawn from the
developmental sequence which proceeded from a
study of marshes to a study of the total wetlands
spectrum. This conclusion is — given the direction of
this sequence — marsh must be the ‘whole’ into which
all the pieces are to be fitted. Hence we find such
anomalies as “bog marshes” (which are fens!), “marsh
exploitation by peat mining,” “meadow” (blanket
bog).
An analysis of the chapters of this book which deal _
specifically with wetland ecosystems indicates that
there has been only a modest attainment of the goal of
expanding the scope of the field of study. About 75%
of the content of these chapters is devoted to a
consideration of marshes, 15% considers open fens,
5% considers a combination of open fens and
meadows, and 5% considers the overall category of
wetlands. Hence, the consideration of marshes pre-
ponderates, and swamps, open and treed bogs, and
treed fens, all substantial portions of the wetlands
spectrum, have not been treated in this volume.
Because primary production processes form the
first section of this book it may be assumed that the
editors assign first priority to this aspect of wetland
ecosystems. On lands which provide essential agri-
cultural and forest crops, emphasis on production 1s
warranted and the tedious, time-intensive activity of
quantitative production evaluations is a worthy
endeavor. On wetlands, however, the emphasis on
THE CANADIAN FIELD-NATURALIST
Vole93
production processes is not justified. If the case for the
conservation of wetlands is to be convincingly
presented to legislators and to the public, conserva-
tionists and scientists should give highest priority to
understanding the role of wetlands in serving as
groundwater recharge areas, as stream-flow regu-
lators (roles which, at best, achieve no more than brief
references in this book), in serving as wildlife habitats
and as an important component of an aesthetically-
pleasing, varied landscape. Accepting this book’s
emphasis on marshes, I must observe I have not
experienced any competition in my annual harvests of
Typha latifolia pollen and a significant harvest of
marsh emergents for cattle fodder is unlikely since it
would seriously conflict with other marsh uses and
because, as indicated in this book, the bulk of the
mineral nutrients of marshes are contained in the
substrata.
I shall temper my predominately negative re-
marks with a few concluding, positive observations.
The Summary and Recommendations sections, which
conclude each of the four parts of the book, provide
good summary statements on the stage of develop-
ment of the art and the high priority projects for future
work on the four ecosystem aspects — primary
production processes, development processes, nutri-
ent dynamics, and management potential.
The contribution of Richardson et al. (“Nutrient
Dynamics of Northern Wetland Ecosystems”) utili-
zes a rational wetlands classification scheme so that
some of the important nutrient and environmental
criteria which characterize the different wetland types
begin to emerge.
The contribution of Sloey et al. (“Management of
Wetlands for Nutrient Assimilation”) is an excellent
dissertation on a topic which ts likely soon to be an
environmental concern. His cautionary statement is
worth quoting for The Canadian Field- Naturalist
readers: “In the past we caused deterioration of the
quality of our surface waters by using them to treat
our wastes. Lest we make the same mistakes in
handling our valuable and diminishing wetlands, it is
mandatory that we carry out long-term, carefully-
monitored experiments at a severely-limited number
of sites. It is important that those conducting these
experiments document changes in the natural system
that could signal future problems.”
A. N. BOISSONNEAU
Ontario Ministry of Natural Resources, Ontario Centre for
Remote Sensing, 880 Bay Street, Toronto, Ontario MS5S
1Z8
1979
ENVIRONMENT
Boreal Ecology
By William O. Pruitt. 1978. Studies in Biology No. 91.
Arnold (Canadian distributor MacMillan, Toronto).
73 pp., illus. Paper $4.80; cloth $10.20.
Boreal ecology is a book of definition and hence one
of introduction. Like most other books in this series it
is designed to keep teachers and students alike
informed of recent developments in biology. It is a
book about snow, and to this end Pruitt maintains
that the . . . “ecological effects of snow cover are so
multi-faceted and all prevading that one might
accurately state that boreal ecology is the study of the
ecology of snow.”
The book deals in generalities and briefly covers the
following aspects of boreal ecology: radiant energy
and light; water, soil and permafrost; bioclimate:
characteristics of boreal vegetation; boreal animals,
complete with a discussion of some of their adapta-
tions to the boreal environment, ecosystems, and food
webs: human utilization of boreal regions: research
methods and procedures peculiar to boreal regions.
Behind Pruitt’s exacting and scientific description
of the boreal forest regions, there is a message. In
essence, it centers upon his belief that continued
misuse of this fragile environment means inevitable
BOOK REVIEWS
D5)
degradation and eventual destruction. Disruption of
the faunal contingent of the boreal forest is well
illustrated in the case of the Alaska Tundra Vole
(Microtus oeconomus). Populations of this vole
have extended their range from normal riverine
locations into the crevices formed from tractor trains,
which through repeated use have exposed the perma-
frost and subjected it to severe melting. Says Pruitt,
“|. . one can only speculate on the possible genetic
results of having a population attenuated to one or
two Microtus wide and perhaps thousands of Micro-
tus long.” It seems to me that we should begin to stop,
look, and listen before unknowingly stepping into
trouble.
The text is well organized, easy to follow and
supported with maps, diagrams, labelled photographs
and tables. The reference list is extensive enough to
serve as the beginning point of a literature search.
It is an excellent piece of work and I recommend it
to interested beginners and professionals.
PAUL A. GRAY
Ontario Ministry of Natural Resources, Box 89, Cochenour,
Ontario POV ILO
The Environmental Impact of Outdoor Recreation
By G. Wall and C. Wright. 1977. Department of Geog-
raphy Publication Series No. 11, University of
Waterloo, Waterloo. x + 69 pp., illus. Paper $5.
This book evolved out of *...a need to synthesize
the research on environmental impact of recreation,
to draw planning and management implications from
the research, and to identify areas of further research.”
The text is organized into nine sections: the
introduction, which outlines the problem, the study,
and the methodology in environmental impact re-
search: geology; soil; vegetation; water quality:
wildlife: air; implications and conclusions, complete
with planning and site management recommenda-
tions; and references.
The impact of outdoor recreation on geology and
air, determined by the authors not to be a major
problem, is given little space in the analysis. Soil is
considered at length; however, most of the literature
comes from the United States and England, and
consequently, in the Canadian context much of the
research has little meaning. Compaction, organic
matter, soil nutrients, temperature, run-off and
drainage changes, erosion, and soil organisms are
considered. The authors suggest that more research is
required on temperature, soil organisms, bacteria,’
and nutrients.
Vegetation, which is usually first to be affected by
physical disturbance, is reportedly well covered in the
literature. It is reviewed in terms of change in
percentage cover and species diversity, plant growth
and age structure, and habitat diversity and mechan-
ical damage. The authors believe that more work is
required because the research to date has“... yielded
conflicting results, especially in terms of compaction,
soil moisture, nutrient levels and soil organisms, and
their associated affects on vegetation species and
growth rates.”
A considerable amount of research into water
quality has been conducted in Canada, especially in
Ontario. This work has been completed primarily out
of concern for the detrimental effects that cottaging,
camping, boating, fishing and other related activities
216
might have on the aquatic ecosystems. Wall and
Wright concentrate their discussion on pathogens,
nutrients and plant growth, dissolved oxygen, and
pollutants. They suggest that research must continue
because “... there is a lack of quantitative analyses of
change in water quality brought about by different
types and intensities of recreational use.”
The wildlife section consists of a discussion on
disturbance, loss and gain of habitats, alteration of
population, and changes in species composition. In
my opinion the authors have unsuccessfully at-
tempted to summarize the impact of recreation on
wildlife in flowchart fashion. They list the immediate
results of intrusion into a wildlife habitat as killing,
disturbance and alteration, which results in adapta-
tion, or migration, and a change in reproduction
levels. The final result is a change in the population
and a different species composition.
It goes without saying that upon intrusion into a
wildlife habitat the first three events occur; however, I
believe that the primary effect of an intrusion is an
alteration of habitat, which in turn causes disturbance
L’inventaire du Capital Nature: méthode de
territoire (3éme approximation)
By M. Jurdant, J. L. Bélair, V. Gerardin, and J. P. Ducruc.
1977. Péches et Environnement Canada, Québec. Ap-
provisionnements et services Canada, Ottawa. 202 pp.,
illus. $7.00, $8.40 (other countries) French.
The purpose of this volume, aimed at those
individuals involved in land management, is to
introduce a method for land classification and
ecological survey. It has taken ten years of field
verification and re-evaluation to perfect this method
and present it in a workable document. Initially the
development of the method was started in 1967 asa
pilot project in the Saguenay/Lac-St.-Jean region of
Quebec; its workability was verified when Jurdant’s
team undertook the ecological survey of 350 000 km?
in the James Bay area.
Chapter | outlines the goal and objectives of the
ecological inventory. The goal is to produce carto-
graphic documents that are characterized by the most
permanent bio-physical parameters of the environ-
ment and hence provide an ecological basis for
planning the integrated management of the resources
of a given territory. They achieve this goal by
producing an ecological map that summarizes the bio-
physical parameters investigated.
In Chapters 2 and 3, the authors discuss the basic
concepts and criteria used in ecological classification.
THE CANADIAN FIELD-NATURALIST
Vol. 93
resulting in productivity alteration, mortality, emi-
gration or immigration, or adaptation. A change in
the population and the species composition then
occurs.
The book is reasonably well organized and comes
complete with graphs, flowcharts, and a reference list.
The composition is weak in a few areas and the
authors have a tendency toward wordiness; for
example, “Even without the intervention of man, the
environment would not be unchanging, but would be
in a perpetual state of flux.” They might have said,
‘Even without the intervention of man, the environ-
ment is naturally dynamic.’
I recommend this book to those unfamiliar with this
subject and interested in developing a basic under-
standing of environmental impact.
PAUL A. GRAY
Ontario Ministry of Natural Resources, Box 89, Cochenour,
Ontario POV ILO
classification et de cartographie écologique du
The basic taxonomic unit for the inventory is the
ecological phase which is defined by the combination
of certain vegetation and soil characteristics (this uses
the concept that vegetation and soil are the best
integrators of variation in biosystems and geosystems
respectively). For the method to work the classifica-
tion criteria must indicate the intrinsic properties and
the actual state of the ecosystem. The criteria must be
measurable and should contribute in some way to
vegetation evolution. Twenty-one criteria are listed
for both terrestrial and aquatic ecosystems.
Chapters 4 to 9 describe in very precise terms the
levels of ecological detail that are observable at
different cartographic scales, the methods of classifi-
cation and cartography, and the methods for analyz-
ing vegetation. An example of what and how to
measure, how to record data at all ecological reference
sites, and how the cartography is interpreted and
applied for management purposes is given. The maps
and annexes demonstrate the workability of the
ecological survey method.
As I have indicated to many of my colleagues, this
book will be an invaluable asset to ecologists,
planners, and managers at all levels of government,
and to private industries and especially to those
involved with the preparation of environmental
impact statements.
1979
The book is replete with original drawings and
caricatures by L. Renaud, that add to the readability
of the prose. In addition, the authors have included
several flow charts depicting the basic concepts of the
ecological survey and outline how a multidisciplinary
team should be organized to carry out a successful
BOOK REVIEWS
2A)
ecological inventory.
GILLES ROBITAILLE
Environment Canada, Laurentian Forest Research Centre,
Box 3800, Ste-Foy, Québec GIV 4C7
Applications de la Télédétection a Etude de la Biosphére
By C. M. and M. C. Girard. 1975. Masson, Paris. 186 pp.,
illus. Paper in French. 96F.
Remote sensing techniques (both instrumentation
and application) have developed very rapidly since
1970. This text, published in 1975, is very scientific in
its approach, is extremely well documented, is clear
and to the point, is logically structured, and is very
particular to details in methodology and definition.
The work examines a wide spectrum of applications
and the in-depth analysis of examples will certainly be
appreciated by the readers. The Girards have or-
ganized this work into four parts, each complete in
itself.
In part one, ‘Remote Sensing,’ the authors present a
discussion of the principles involved. They describe
the characteristics and the energy sources of the
electromagnetic spectrum, the influences of the
atmosphere, and explain the mechanisms involved for
the peculiar behavior of vegetation, soil, water, ice
and snow in different spectral ranges such as the
visible and near infrared, middle and far infrared, and
the very high frequencies. Both photo and non-
photographic instrumentation is described. Data
treatment analysis, i.e., visualization, increasing con-
trast, and information selection, as well as factors that
influence the utilization of data are given.
Part Two is entitled ‘Methods’ with the emphasis
being placed on methods of photo and image
interpretation.
Sixty percent of the book (Part Three) is devoted to
‘Interpretation.’ It is a detailed account applied to
vegetation, pedology, agronomy, and to anthropo-
genic influences. Text examples are often related to
actual photographs. The interpretation of vegetation
is looked at from the point of view of species
identification, recognition of vegetation assemblages,
edaphic conditions, and succession of plant com-
munities. The pedology section looks at the ident-
ification of soil characters that are both readily and
not readily discernable. Examples are also given. The
interpretation of agricultural lands is also treated. The
authors give information on crop identification, soil
modification, and touch on the problem of agronomic
cartography. They also expand on the utilization of
remote sensing for pest control. Anthropogenic
influences on the environment are increasing. The
artificial interferences by man that lead to urban-
ization are not treated. What the authors do look at is
the interpretation of influences on the natural
environment and rural modifications.
In Part Four an example of how high-altitude data
(35 km high from balloons) are utilized in inter-
pretation and thematic cartography (..e., geology, soil
utilization) is given. This part concludes with a flow
chart on data treatment by computers.
During the three years since publication, many
innovations have occurred in the field of remote
sensing. This text remains a valuable source of
material for those interested in acquiring a basic
applications reference. It is recommended for use at
the university level and by current practitioners.
GILLES ROBITAILLE
Environment Canada, Laurentian Forest Research Centre,
Box 3800, Ste-Foy, Québec GIV 4C7
Wildness Is All Around Us: notes of an urban naturalist
By Eugene Kinkead. 1978. Dutton, (Canadian distributor
Clarke Irwin, Toronto). 178 pp., illus. $12.50.
Wildness Is All Around Us is an informative and
entertaining book that offers a variety of insights into
some items of natural history, plus many entrancing
digressions. For example, the essay on the coyote
(pronounced ky oat) describes not only its dis-
tribution, paleontological and recent history, appear-
ance, behavior, relationship with man, and repro-
duction with other canids, but also includes lengthy
asides on endangered species (of which the coyote 1s
certainly not one), on sheep and sheep-ranching, on
218
wolves, and on the principles of heredity. But there is
almost nothing about coyotes in cities, despite the
subtitle of the book.
Kinkead also includes a long essay on the House
Sparrow, a fascinating history of this species’ intro-
duction to North America (Manhattan) in and after
1848 to reduce the measuring-worm population, (even
though anti-sparrow organizations existed in England
as early as 1744), and its subsequent conquest by
1880 of much of the continent. Until that time these
widely-acclaimed birds were shipped freely about the
country to eradicate noxious insect pests. After that
time, however, the sparrow was seen rather as “an evil
of monstrous proportions.” Sparrows molested over
70 species of native birds; ate fruits, vegetables, grain
crops, and urban plantings; and around dwellings
were dirty, noisy, and blatantly promiscuous. The
New York Times predicted in 1887 that once
Americans realized how tasty sparrows roasted with
bacon were, their numbers would be decimated by
cooks, but this did not happen. Instead sparrow
populations continued to expand until about 1910,
when the increase in the automobile and consequent
decrease of horses and the grain they ate undercut the
sparrows’ food supply, especially in winter and in
cities. Since then sparrow populations, although still
numerous, have stayed within more reasonable
bounds.
The other chapters in this book are really vignettes
rather than essays. The first describes the pande-
monium which took place one spring when a Steller’s
OTHER
THE CANADIAN FIELD-NATURALIST
Woln9s
Eider from Siberia was spotted off the Boston South
Shore. The area was soon swamped with birders
wanting to increase their Life List, alerted by a
message taped for the “Voice of Audubon” Boston
telephone number. The second is about a rare birch
tree, Betula uber, rediscovered with the help of the
author after 60 years beside a creek in the Appala-
chians of Virginia. The geological history and the
wildflowers of Central Park in New York City are the
subjects of the other two chapters.
Although Eugene Kinkead is primarily a writer
rather than a biologist, he includes in this book a
wealth of biological information. He is not correct,
however, in saying that no person in North America
has been attacked by a healthy wolf. In northern
Ontario in 1942 a wolf knocked a railman off his
handear, and, while canoeing in 1893 through the
barren lands of Canada, J.B. Tyrrell’s party was
beseiged by wolves. Similar attacks have occurred in
Eurasia. I think also few would agree with the dust-
cover statement that the fox is threatened with
extinction.
This well-written book with its few, evocative
sketches uses a format which could profitably be a
model for many would-be authors writing about
nature. | recommend it highly to both laypeople and
professional biologists.
ANNE INNIS DAGG
Box 747, Waterloo, Ontario N2J 4C2
Field Photography: beginning and advanced techniques
By Alfred A. Blaker. 1976. Freeman, San Francisco.
(Canadian distributor Oxford University Press,
Toronto). 450 pp., 55 black and white plates, 12 color
plates, plus 41 pp. field book. $19.95.
This book is a very comprehensive treatise on the
subject of field photography. It follows the same
format and style as the author’s previous publication,
Photography for Scientific Publication: a handbook
(Freeman, San Francisco, 1965), which presented a
synthesis of laboratory photographic technique. The
purpose of this publication is instructional; the author
states in the preface that, “this book will represent a
self-teaching aid to help interested persons improve
the quality of photographs they may need.” The text
accomplishes this purpose.
The book is organized into three sections: Back-
ground, Basic Photography, and Field Photography
Techniques. There are also three appendices contain-
ing film speed data as well as development and
copying information. A supplemental pamphlet.sum-
marizes mathematical tables and formulae in a form
suitable for field use.
The background section introduces the reader to
scientific field photography. It discusses merits of
photography, photographic training, and offers ad-
vice on photographic operations. This information is
valuable to persons currently involved in professional
photographic capacities. The author continues this
section with a discussion on the tools of photography.
He accurately and extensively describes cameras, film,
1979
development materials, and essential accessories used
in modern photography. This discussion has two
minor, unavoidable problems: personal bias and some
outdated information.
The second part of the text, basic photography, is
most informative to the incipient nature photog-
rapher. An initial short chapter describes considera-
tions related to composition of photographs, oras the
author puts it, “seeing a picture.” This section is very
informative, and is highly recommended for both
inexperienced and veteran photographers.
Blaker next presents explanations concerning basic
topics of photography such as exposure, photo-
graphic filters, and darkroom procedures. The data
given here are detailed and it is recommended that the
reader be concerned only with problems of his/her
particular interest. Although the writing style here is
clear enough, there is some difficulty in grasping the
extensive reference material in one quick reading. If a
simplistic discussion of these fundamental topics is
required one should seek elsewhere.
The final portion of the text deals specifically with
field techniques. Blaker commences this section with
an appeal to field workers to maintain ethical
ecological considerations while endeavoring to cap-
ture a picture. This is sound advice that all nature
photographers should attend. A short section on
climatic problems in photography and proper equip-
ment preparation for field work, rounds out this
interesting division of text.
The remaining chapters of this section contain
information on essential topics of nature photog-
raphy: close-up, long-range (telephoto), and wide-
angle photography. The author begins with close-up
(macro) imaging. He explains equipment, back-
ground theory (of focal length changes, etc.) and
comments on special problem areas. This information
NEW TITLES
Zoology
Avian breeding cycles. 1977. By R. K. Murton and N. J.
Westwood. Clarendon (Oxford University Press, New
York). xiv + 594 pp., illus. US$48.
The behavior of fish and other aquatic animals. 1978.
Edited by D. I. Mostofsky. Academic, New York. xiv + 394
pp., illus. US$27.50.
*Behavioural ecology: an evolutionary approach. 1978.
Edited by John R. Krebs and Nicholas B. Davies. Black (US
and Canadian distributor Sinaver, Sunderland, Maryland).
512 pp., illus. Cloth US$34; paper US$17.50.
BOOK REVIEWS
AIS)
is very useful, especially the data on close-up
photography with flash equipment.
The chapter on telephoto work is unfortunately not
as helpful. Some advice here is questionable; for
example, the point is made that teleconverters give as
good (or better) results for a portion of the cost of
telephoto lenses. A pertinent point overlooked is that
the image one obtains 1s directly related to the quality
of the lens with which it 1s captured. Blaker does not
explain that teleconverters of sufficient optical quality
are not inexpensive. A telephoto lens with its added
versatility may on occasion be a better choice of
equipment.
The next chapter gives a brief description of wide-
angle photography. Although little emphasis is placed
on landscape photography, the remainder of the
chapter is complete.
The last two chapters deal with stereo photography,
which is not an essential element in most field work.
But this section presents interesting information that
may invoke novel methodology in future field
photography.
Field Photography 1s a book that offers something
for every nature photographer. The more experienced
will want this text for use of its inexhaustive reference
data and extensive bibliography. Beginners will find
the material on basic photographic procedures very
worthwhile, and advice on problem areas invaluable.
All will appreciate its wide scope, logical presentation,
and picture quality. A word of caution: this publica-
tion is designed as a reference text and is written
accordingly.
ALAN J. KENNEDY
11416-50 Avenue, Edmonton, Alberta T6H 0J3
Bird flight. 1978. By Georg Ruppell. Translated from the
German edition (Munich, 1975). Van Nostrand Reinhold,
New York. 192 pp., illus. US$18.95.
Birds as builders. 1977. By P. Goodfellow. Arco, New York.
168 pp., illus. US$15.95.
*Birds of man’s world. 1978. By D. Goodwin. Cornell
University Press, Ithaca. vi + 183 pp., illus. US$10.45.
A birdwatcher’s adventures in tropical America. 1977. By
A. F. Skutch. University of Texas Press, Austin. xii + 327
pp., illus. US$13.95.
220
*A bird watcher’s guide to the eastern United States. 1978. By
Alice M. Geffen. Barron’s Education Series (Canadian
distributor Burns and MacEachern, Toronto). 384 pp., illus.
US$15.95.
t+Ducks, geese and swans of the world. 1978. By Paul A.
Johnsgard. University of Nebraska Press, Lincoln.
xxl + 404 pp., illus. US$35.
East African mammals. An atlas of evolution in Africa.
Volume 3, part A (Carnivores). 1977. By J. Kingdon.
Academic, New York. vill + 476 pp., illus. US$74.25.
Eels: a natural and unnatural history. 1978. By C. Moriarty.
Universe, New York. 192 pp., illus. US$1S.
*The Gannet. 1978. By Bryan Nelson. Butes, Vermillion,
South Dakota. 336 pp., illus. US$25.
Granivorous birds in ecosystems. 1977. Edited by J.
Pinowski and S. C. Kendeigh. Cambridge University Press,
New York. 451 pp., illus. US$47.50.
A guide to fishes of the temperate Atlantic coast. 1977. By
M. J. Ursin. Dutton, New York. xx + 268 pp., illus. Cloth
US$10.95; paper US$5.95.
Insect life. 1977. By M. Tweedie. Collins, London. 192 pp..
illus. US$8.95.
{Migratory game bird hunters and hunting in Canada. 1978.
Edited by H. Boyd and G. H. Finney. Canadian Wildlife
Service Report Series Number 43. Supply and Services
Canada, Hull. 125 pp., illus. $7.50 in Canada; $9 elsewhere.
Octopus. Physiology and behaviour of an advanced inverte-
brate. 1978. By M. J. Wells. Chapman and Hall, London
and Halsted (Wiley), New York. xiv + 418 pp., illus.
US$42.50.
+The order Microsauria. 1978. By R.L. Carroll and P.
Gaskill. Memoirs Volume 126. American Philosophical
Society, Philadelphia. 211 pp., illus. Paper US$15.
Parental behavior in birds. 1977. Edited by R. Silver.
Dowden, Hutchinsonand Ross, Stroudsburg, Pennsylvania.
xvill + 436 pp., illus. US$24.50.
Quantitative ethology. 1978. Edited by P. W. Colgan. Wiley-
Interscience, New York. xvi + 364 pp., illus. US$25.
Readings in sociobiology. 1978. Edited by T. H. Clutton-
Brock and P. H. Harvey. Freeman, San Francisco. x + 394
pp., illus. Cloth US$18; paper US$9.
Songbirds of the eastern and central states. 1977. By T. L.
Rising. Tundra (Scribner’s), Plattsburgh, New York. viii +
48 pp. US$3.95.
Studer’s popular ornithology: the birds of North America.
1977. Edited by J. H. Studer. Original publication 1881.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Harrison House/ Barre (Crown), New York. x + 301 pp.,
illus. US$39.95.
Sulidae. Gannets and boobies. 1978. By J.B. Nelson.
Aberdeen University Study Series, Number 154. Oxford
University Press, New York. xii + 1012 pp., illus. US$98.
Transactions of the XIII International Congress of Game
Biologists. 1977. Proceedings of a conference in Atlanta,
Georgia 11-15 March, 1977. Wildlife Society, Washington.
538 pp. Paper US$10.
*Vanishing birds. Their natural history and conservation.
1978. By Tim Halliday. Holt, Reinhart, and Winston,
Toronto. US$16.95.
Ways of wildlife. 1977. Edited by E. Horwitz. Wildlife
Society, Washington. 157 pp. Paper US$2.95.
Botany
Dangerous plants. 1977. By John Tampion. David and
Charles (Canadian distributor Douglas, David and Charles,
Vancouver). 176 pp., illus. $12.50.
Darwin and his flowers: the key to natural selection. 1977. By
M. Allan. Taplinger, New York. 318 pp., illus. US$14.50.
Fenland. Its ancient past and uncertain future. 1978. By H.
Godwin. Cambridge University Press, New York. 203 pp..,
illus. US$18.95.
The flora of Canada: part 1 — general survey; part 2 —
Pteridophyta, Gymnospermae, Monocotyledoneae; part 3
— Dicotyledoneae (Saururaceae to Violaceae). 1978. By
H. J. Scoggan. Publications in Botany Numbers 7-1, 7-2,
and 7-3. National Museums of Canada, Ottawa. 1115 pp. $89
for 3-volume series in Canada: $106.80 elsewhere.
An illustrated history of herbals. 1977. By F. J. Anderson.
Columbia University Press, New York. xvi + 270 pp., illus.
US$16.95.
Important forest trees of the United States. 1978. By U.S.
Forestry Service. Public Documents Distribution Center,
Pueblo, Colorado. 70 pp., illus. US$2.10 plus 25% foreign’
handling.
Introduction to world vegetation. 1978. By A. S. Collinson.
Allen and Unwin, Winchester, Massachusetts. 192 pp., illus.
Paper US$8.75; cloth US$16.95.
Manual of woody landscape plants: their identification,
ornamental characteristics, culture, propagation and uses.
1978. Stripes, Champaign, Illinois. 536 pp., illus. $15.
The moss flora of Britain and Ireland. 1978. By A. J. E.
Smith. Cambridge University Press, New York. C 900 pp..,
illus. US$59.50.
1979
Orchid biology: reviews and perspectives, I. 1977. Edited by
J. Arditti. Comstock (Cornell University Press), Ithaca. 312
pp., illus. US$29.50.
+Physiology and biochemistry of seeds in relation to
germination. Volume 1, development, germination and
growth. 1978. By J. D. Bewley and M. Black. Springer-
Verlag, New York. xi + 306 pp., illus. US$45.
River plants. 1978. By S. M. Haslam. Illustrated by P. M.
Wolseley. Cambridge University Press, New York. C 350
pp., illus. Cloth US$62.50; paper US$14.95.
{Tropical trees and forests. An architectural analysis. 1978.
By F. Hallé, R. A. A. Oldeman, and P. B. Tomlinson.
Springer-Verlag, New York. xvi + 441 pp., illus. US$62.50.
Tropical trees as living systems. 1978. Edited by P. B.
Tomlinson and M. H. Zimmerman. Cambridge University
Press, New York. C 624 pp., illus. US$49.95.
*Vascular plant families. 1977. By James Payne Smith, Jr.
Mad River Press, Eureka, California. 320 pp. US$7.85.
+ Wild coffee and tea substitutes of Canada. 1978. By Adam
Szezawinksi and Nancy Turner. Edible Wild Plants of
Canada, Number 2. National Museum of Natural Sciences,
Ottawa. I11 pp., illus. $6.95.
Environment
Ecology and environmental planning. 1978. By J. M. and
M. A. Edington. Chapman and Hall, London and Halsted
(Wiley), New York. viii + 246 pp., illus. US$22.50.
Ecology field glossary: a naturalist’s vocabulary. 1977. By
W. H. Lewis. Greenwood, Westport, Connecticut. x11 + 152
pp.. illus. US$1S.
*The ecology of North America. 1978. By V. E. Shelford.
First paperback edition (original publication 1963). Uni-
versity of Illinois Press, Urbana. 610 pp., illus. No price
given.
Enjoying nature with your family. 1977. By M. Chinery.
Crown, New York. 192 pp., illus. US$12.95.
Environmental assessment law in Canada. 1978. By Paul D.
Edmund. Edmund-Montgomery, Toronto. xx + 380 pp.
$30.
tEnvironmental chemistry and cycling processes. 1978.
Edited by D. C. Adriano and |. L. Brisbin, Jr. Proceedings
of a symposium, Augusta, Georgia, April 28-May 1, 1976.
U.S. Department of Energy Symposium Series Number 45.
National Technical Information Service, Springfield, Vir-
ginia. 943 pp., illus. Paper US$I5.
Environmental issues. 1978. Edited by M. W. Holdgate and
G. F. White. SCOPE 10. Wiley, Toronto. 242 pp. Paper $23.
BOOK REVIEWS
Up)
Environment and society: an introductory analysis. 1977. By
B. Harvey and J.D. Hallett. MIT Press, Cambridge.
x + 163 pp., illus. Paper US$6.95.
The limits of altruism: an ecologist’s view of survival. 1977.
By Garrett Hardin. Indiana University Press, Bloomington.
vi + 154 pp., illus. US$10.
Nature/science annual, 1978 edition. 1977. By the editors of
Time-Life Books. Time-Life, Alexandria, Virginia. 192 pp.,
illus. US$7.95.
Nature’s economy: the roots of ecology. 1977. By D.
Worster. Sierra Club, San Francisco. xii + 404 pp. US$15.
Pacific seashores: a guide to intertidal ecology. 1978. By
Thomas Carefoot. University of Washington Press, Seattle.
208 pp., illus. Paper US$12.95.
Principles of pollination ecology. 1978. By K. Faegriand L.
van der Pil. Third revised edition. Pergamon, New York.
242 pp., illus. Cloth US$25; paper US$15.
Pyramids of life: illuminations of nature’s fearful symmetry.
1977. by J. Reader and H. Croze. Lippincott, Philadelphia.
222 pp., illus. US$12.95.
Risk assessment of environmental hazard. 1978. By R. W.
Kates. SCOPE 8. Wiley, Toronto. 160 pp. Paper $13.35.
A Sierra Club naturalist’s guide to the deserts of the
southwest. 1977. By P. and L. Larson. Sierra Club, San
Francisco (Canadian distributor Wiley, Toronto). 288 pp..,
illus. Cloth US$9.59; paper US$5.95 (Cdn. $7.95 paper).
The thin edge: coast and man in crisis. 1978. By A. W.
Simon. Harper and Row, New York. xii + 180 pp. US$10.
The underwater wilderness: life round the great reefs. 1977.
By C. Roessler. Chanticleer (Dutton), New York. 319 pp.,
illus. US$35.
*Vermilion Lakes Banff National Park: an introductory ©
study. 1978. By Bow Valley Naturalists, Banff, Alberta.
iv + 68 pp., illus. Paper $3.
Miscellaneous
Astronomy now. 1978. By J.M. Pasarhoff. Saunders,
Philadelphia. xvi + 400 pp., illus. Paper US$1I1.95.
Astronomy, the cosmic journey. 1978. By W.K. Hart-
mann. Wadsworth, Belmont, California. x11 + 536 pp., illus.
US$16.95.
Bibliography of published and unpublished literature on the
Hudson Bay Lowland. 1978. By S. E. Haworth, D. W.
Cowell, and R. A. Sims. Report O-X-273. Great Lakes
Forest Research Centre, Sault Ste. Marie. 270 pp. Free.
Bio-babel. Can we survive the new biology? 1978. Edited by
Allen R. Utke. John Knox Press, Atlanta. 248 pp. US$11.95.
22D
Biométrie et écologie. 1978. By J.M. Legay and R.
Tomassone. Societé frangaise de Biométrie, Jouy-en-Josas,
France. xi + 388 pp., illus. US$8.
The faces of the Great Lakes. 1977. By J. Ela. Sierra Club,
San Francisco. 192 pp., illus. US$24.50.
Hunger on planet earth. 1977. By J. Archer. Crowell, New
York. vill + 216 pp., illus. US$7.95.
Land/water classification. 1978. By D.M. Welch. Eco-
logical Land Classification Series Number 5. Supply and
Services Canada, Hull. x +54 pp., illus. Paper $4.50 in
Canada; $5.40 elsewhere.
+ Last of the naturalists: the career of C. Hart Merriam. 1977.
By K.B. Stirling. Revised edition. Natural Sciences in
America Collection. Arno, New York. 472 pp., illus. US$23.
*Lexique anglais-francais, termes techniques a lusage des
biologistes. 1978. By Jean Vaillancourt. Editions de ?Uni-
versité d’Ottawa, Ottawa. 427 pp. $12.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Nature to be commanded. Earth-science maps applied to
land and water management. 1978. By U.S. Geological
Survey. Public Documents Distribution Centre, Pueblo,
Colorado. 96 pp., illus. US$6.25 plus 25% foreign handling.
Simulation modelling of environmental problems. 1978.
Edited by F. N. Frenkiel, D. W. Taylor, and D. W. Goodall.
SCOPE 9. Wiley, Toronto. 128 pp. Paper $13.35.
Sun/earth. How to use solar and climatic energies. 1978.
By Richard C. Crowther. Scribner, New York. vil + 232 pp.,
illus. Paper US$8.95.
The world of Roger Tory Peterson: an authorized biog-
raphy. 1977. By J. C. Devlin and G. Naismith. Illustrations
and paintings by Roger Tory Peterson. Quadrangle (New
York Times), New York. xxii + 266 pp., illus. US$14.95.
+Available for review
*Assigned for review
The Ottawa Field-Naturalists’ Club
Minutes of the Ninety-Ninth Annual Business Meeting of The Ottawa Field-Naturalists’ Club
The 99th Annual Business Meeting of The Ottawa
Field-Naturalists’ Club was held in the auditorium of
the National Museum of Natural Sciences, on 10
January 1978. The President, R. A. Foxall, called the
meeting to order at 8:17 p.m., with a quorum of 26
persons present (final total, 27). The Recording
Secretary read the minutes of the 98th Annual
Meeting, which were approved on motion (by K.
Strang, 2nd, H. Thomson).
R. Foxall referred to business arising from the
minutes. The Council had directed the Publications
Committee to consider the status of The Shrike as a
Club publication; the Committee has recommended
that The Shrike be considered a publication of The
Ottawa Field-Naturalists’ Club, a recommendation
accepted by the Council. As recommended at the last
Annual Meeting, the Conservation Committee had
considered the matter of the necessity for approval by
Council of all reports and briefs; and had reeommend-
ed that all significant submissions, containing policy
statements or contentious issues, should be cleared by
the President or his designate. A General Meeting of
the Club, on Centennial planning, had been held in
June 1977, to inform the membership of the projects
being undertaken, and to get ideas of the interests of
the members.
In the absence of the Treasurer, the Club’s
accountant, M. Brigham, was called upon to present
the 1977 financial report. Unfortunately, this had not
been audited, owing in part to the current prevalence
of the ‘flu’. R. Taylor, the Chairman of the Finance
Committee, explained that the change of the financial
year, to end at the end of September, was to relieve the
pressure on the Treasurer’s Assistant and the Auditors
to produce the statement in time for the Annual
Business Meeting; closing the books in December had
always been difficult, particularly with membership
dues still coming in at that time. For this reason, this
year’s statement covers only nine months. Taylor also
explained that the apparent large profit to the Club
was due to the lack of allocation of funds to The
Canadian Field- Naturalist. Excluding ‘windfall’ items
such as life memberships, which cannot be budgeted,
the Club had actually operated at a slight loss,
justifying the increase in membership dues for 1978.
The Canadian Field- Naturalist profit was due mainly
to the success of the raptor issue; the grant request to
the National Research Council for 1978 has been
reduced accordingly.
R. Foxall raised the point that the Auditors,
approved at the last Annual Meeting, were G. J.
Wasteneys and D. Potter; since that time, D. Potter
has changed his job and will not have time to do the
audit before the end of February. Wasteneys is still
available, but the Constitution requires two auditors.
C. Gruchy moved (2nd, E. Dickson) that the
responsibility for finding a second auditor for the
financial year ending September 30, 1977 be delegated
by the Annual Business Meeting to the President of
The Ottawa Field-Naturalists’ Club; this was approv-
ed unanimously. Gruchy then questioned the need for
the Club to have two auditors: M. Brigham agreed
that one person could do the job. It was recommended
that this matter should be examined by the Council to
see whether a change to the Constitution may be in
order.
R. Taylor moved (2nd, C. Gruchy) adoption of the
interim, unaudited, financial statement covering |
January 1977 to 30 September 1977: the motion was
approved.
The Annual Report of Council, to be published in
The Canadian Field-Naturalist, was read by R.
Foxall. A. Erskine pointed out that there were
deficiencies in, rather than lack of, knowledge of our
area’s natural history: and Todd suggested that the
Pink Road feeder should in some way be identified as
the Lucerne feeder mentioned in the previous report.
Acceptance of the report, as amended, was moved by
S. Hamill, 2nd, C. Gilliatt, and approved.
M. Brigham moved a vote of thanks (2nd, J.
Harrison) to all the volunteers who look after the
Club’s bird feeders: George McGee and Bill Holland,
Mr. and Mrs. Hugh Munro and W. Earl Godfrey,
Jean Hastie, and John Dubois. Motion approved
unanimously.
Foxall then called on C. Gruchy, as Chairman of |
the Nominating Committee, to present the slate.
Gruchy apologized for not having published a request
for nominations, but said that despite this, one
nomination had been received from the membership
at large. He then presented the slate:
President: R. A. Foxall; Vice-President: R. Taylor;
Treasurer: B. Henson; Recording Secretary: D.
Laubitz; Corresponding Secretary: S. Armstrong:
additional members of Council: E. Beaubien, C.
Beddoe, W. Cody, J. Diceman, E. Dickson, A. Dugal,
C. Gilliatt, C. Gruchy, P. Hall, J. Harrison, V. Hume,
H. MacKenzie, J. Murray, M. Ney, G. Patenaude, K.
Strang, S. Teeple, E. Todd.
On motion (by C. Gruchy, 2nd, M. Ney) all of those
nominated were declared elected. R. Foxall, on behalf
of the Club, gave thanks to those Council members
who were retiring: Tony Erskine, Gavin Nicholson,
Gerry Oyen, Pamela Sims, and Stan van Zyll de Jong.
223
224
The Auditors for 1977-1978, G. J. Wasteneys and
M. Brigham, were approved on motion(by R. Taylor,
2nd, E. Todd).
Foxall then reported that plans for some of the
Centennial projects are proceeding well, but that
despite many appeals there have been very few
volunteers, and some of the planned activities may
have to be dropped. It is felt that the Club deserves a
good Centennial celebration, but perhaps our ideas
have been too ambitious. We will also need many
volunteers to do small jobs during 1978. E. Dickson
asked whether the list of people who have offered their
skills to help the Club had been consulted by the
Centennial Planning Committee; D. Laubitz answer-
ed that it had been consulted when specific skills had
been needed. M. Brigham raised the point that some
of the proposed Centennial projects concern saleable
items: that the cost of these items may be about equal
to the amount for which they can be sold; and queried
whether consideration had been given to the need for
advertising to promote sales. R. Foxall replied that
THE CANADIAN FIELD-NATURALIST
Vol. 93
the costing of items for sale is indeed a difficult task:
and that advertising was being considered an impor-
tant adjunct to selling Centennial items. He emphasiz-
ed that 1978 will be a very busy year for the Council
and the Club.
J. Harrison moved (2nd, A. Hanes and E. Dickson)
a motion of thanks to Harry Thomson for his long and
dedicated service to the Club, as Business Manager of
Trail & Landscape, from which position he is now
retiring after I] years. The motion was warmly
approved by all present.
E. Todd thanked the President, on behalf of the
Club, for carrying on the business of the Club during
1977, a job which Todd knew by experience to be a
frequently thankless one.
Adjournment of the meeting (on motion by B.
Henson, 2nd, K. Strang) was at 9:35 p.m.
Following the refreshments, color slides were
shown by J. Harrison, M. Ney, and D. Black.
D. R. LAuBITZ, Recording Secretary
Report of Council of The Ottawa Field-Naturalists’ Club
The consideration and support of Centennial
projects has taken up a large proportion of Council’s
time during 1978. One result of the Centennial
program is that we have been reminded that our
members may have in their possession valuable
archival materials referring to The Ottawa Field-
Naturalists’ Club. Club members are therefore re-
minded that any relevant materials for which they no
longer have any use will be gladly received by the
Public Archives.
For the Centennial Steering Committee,1978 was a
period of major activity, leading to Council approval
of 14 of the 33 Centennial projects that had been
proposed. The Club pin, designed by W. Rath, will go
on sale at the Centennial Banquet, to be held on May
19. An exhibition, ‘100 years of looking at nature,’ will
open on May 18 as the preliminary to a weekend of
seminars, workshops, and field trips on natural
history topics. In September, we shall combine a day
of field trips with a Club picnic. Hasty-notes, designed
by E. Dickson, and the Centennial Calendar, featur-
ing works by Club artists, have already been produc-
ed; being prepared for publication are a reissue of
John Macoun’s Autobiography; ‘Orchids in the
Ottawa District’ by A. and J. Reddoch; Cumulative
Indices to Transactions of The Ottawa Field-Natural-
ists’ Club and The Ottawa Naturalist compiled by J.
Gillett; and a revised ‘Checklist of Birds of the Ottawa
Area’ by B. Barrett. A stereo recording of * Nature
Sounds in the Ottawa Area’ by F. M. Brigham is close
to completion. Two projects still in the early stages of
preparation are ‘Birds in the Ottawa Area 1979, a
record of bird sightings and abundance in Centennial
year to be published in 1980; and the Macoun Club’s
natural history ‘Trail and Guide, which was held up
owing to a forced change of location.
(Hue MacKenzie)
The Centennial Ways & Means Committee was set
up to assess Centennial projects’ costs; to determine
the availability of The Ottawa Field-Naturalists’ Club
funds to meet these costs: and to recommend ways of
raising additional funds should they be required.
Total project costs are now estimated to be approxim-
ately $40,500, with recovery through sales of about
$18,400, giving an estimated net outlay of $22,100.
Moneys available through Club funds are about
$50,000: no action to obtain additional funds from
outside the Club was considered necessary.
(C. Gilhatt)
Finance Committee: The 1977-1978 budget was
adopted by Council early in the year. With the
financial year ending in September, the books were
presented to the auditors on time, and the 1978-1979
budget was presented to and approved by Council at
its December meeting. Ivy Hewis resigned as Assistant
1979
to the Treasurer early in 1978: the services of Lois
Cody were engaged for this position temporarily, and
the position was advertised to the Club membership.
Of the three applicants, Mrs. Cody was judged to be
the most suitable, and Council approved her appoint-
ment on a continuing basis. The committee recom- .
mends that the constitutional requirement of two
auditors be changed to one; it also recommends that
one person be appointed to be responsible for the
inventory of sales of all items excluding back issues of
Club journals.
(R. Taylor)
The Membership Committee reported that the total
Club membership now stands at 1150 compared with
1163 in 1977; the membership structure is given in the
table below. The Club lost a well known naturalist and
Honorary Member in January 1978, when Hoyes
Lloyd died. At the Annual Dinner in April, W. K. W.
Baldwin was presented with an Honorary Member-
ship. Marsh Ney, Chairman of the committee,
resigned in August when he moved to Victoria, B.C.
(F. E. Goodspeed)
The Publications Committee reported that, since
the last report, The Canadian Field-Naturalist was
published in four issues, Volume 91(4) and Volume
92(1, 2, 3), with a total of 430 pages containing 37
articles, 46 notes, 37 book reviews, and a listing of 266
new natural history book titles. The publication was
supported in part by a grant of $2200 from the
National Research Council, which is very gratefully
acknowledged. Trail & Landscape was published in
five issues for a total of 152 pages of articles and
information of interest to local members. Marc
Forget, the Business Manager, has had to resign; we
thank him for his contribution to Trail & Landscape.
The Shrike, our local bird-watchers newsletter, was
published in eight issues containing 55 pages. The
committee recommended amendments to By-laws 6
and 9; submitted a grant application for $8000 to the
Natural Sciences and Engineering Research Council
to assist the publication of the indices to the
THE OTTAWA FIELD-NATURALISTS’ CLUB
225
Transactions of The Ottawa Field-Naturalists’ Club
and The Ottawa Naturalist. An analysis of the
authorship of articles published in The Canadian
Field-Naturalist was carried out at the request of
Council. Based on a sample of issues published
between 1950 and 1977, our study indicated that there
had been a steady decline in the number of papers
published by authors not identified with an institu-
tional address. We emphasize The Canadian Field-
Naturalist’s status as a major national, Canadian
publication open to all workers in the field of natural
history; advice and guidance are available to any
author requiring assistance in documenting research
results for publication.
(J. K. Strang)
During 1978 the Excursions and Lectures Commit-
tee organized 10 monthly meetings, with an average
attendance of 30, in the auditorium of the National
Museum of Natural Sciences; three photographic
workshops; the Annual Dinner, with Clarence Tillen-
lus, artist and conservationist, as speaker: and
excursions covering general topics (14), birds (17),
and flowers, butterflies, geology, fish, mushrooms,
and microscopic life (one of each). The bus excursions
are still popular and will be continued: and the use of
younger members as assistant field trip leaders has
been successful both as help to leaders and for training
future leaders. Jeff Harrison, Chairman of the
committee, resigned in May when he moved to
Newfoundland.
(Jeanette Dean)
The Conservation Committee spent most of its time
in 1978 studying and responding to Conservation
Lands in Ottawa-Carleton and the subsequent re-
visions to this study. Proposed changes to the original
(1974) Ottawa-Carleton Regional Official Plan in-
cluded numerous boundary changes and six deletions ~
of Conservation Areas, due in part to pressure from
affected landowners. Asa result of three major reports
prepared by the committee, some of the revised
boundaries were expanded, and two of the deleted
Canadian Canadian
Membership (local) (other) USA Foreign Totals
Individual 454 (438) 321 (342) 102 (106) 3 (5) 880 (891)
Family 198 (207) 26 (22) 1 (2) 2 (1) 227 (232)
Sustaining 9 (10) nil (3) nil nil 9 (13)
Life 13 (9) 7 (6) 3} (1) 2 (2) 25 (18)
Honorary 5 (5) 4 (4) nil nil 9 (9)
Totals 679 (669) 358 (377) 106 (109) 7 (8) 1150 (1163)
Changes +10 -19 -3 —] —13
Figures in brackets represent 1977.
226
areas were proposed for reinstatement. During the
year it became apparent that, despite The Ottawa
Field-Naturalists’ Club representation on advisory
committees and direct communication with planning
staff and politicians, without input by individual
members of the Club the main viewpoint represented
in response to the planning report would be that of
those who were vehemently opposed to conservation
zoning within the Region, as expressed at public
meetings. Letters giving background to the issues were
therefore sent to all local members, requesting them to
write to their local politicians on the issues; there was a
10% response, which was much appreciated. Eleven
field trips, to collect data, were held during the year,
and it is clear that continued activity to try to influence
planners and politicians will be needed for many
months to come. How the committee’s work will
affect the status of conservation in the region is not yet
known.
(Ewen C. D. Todd)
The Macoun Field Club Committee reported that
there are 65 members in the club in this their 30th year.
During 1978 consideration was given to publicizing
the club, but it was decided that, rather than advertise,
information about the club would be distributed to
organizations where it would be readily available to
those seeking such a club; local schools were also
informed of the club’s activities, and the members of
the Senior group offered to speak to schools, and to
lead Club activities, etc. A donation received in
memory of John Bird, from Mrs. Furness Thompson,
of Pennsylvania was used to buy a pair of binoculars
for the club.
(S. Gray)
The Education and Publicity Committee was
responsible for the acquisition and installation of a
THE CANADIAN FIELD-NATURALIST
Vols
Club phone in 1978, with the number to be listed in the
local directory. Other activities included providing
judges for the Ottawa Science Fair, and $100 in prizes;
involvement in the mail campaign for conservation
support; requests for volunteers for weekday activi-
ties: and some advertising of monthly meetings in the
local papers. A special field trip and scavenger hunt
was organized in September for Club members and
their children; it is difficult to tell whether the idea was
well received as it was raining and no one came.
(Elisabeth Beaubien)
The Council has also been concerned with im-
proving relationships both inside and outside the
Club. There has been considerable discussion of the
sort of relationship The Ottawa Field-Naturalists’
Club should have with the Federation of Ontario
Naturalists, and it was concluded that The Ottawa
Field-Naturalists’ Club should have Federated Club
status; the election of Courtney Gilliatt to the
Federation of Ontario Naturalists’ Board of Directors
was welcomed. Long-term discussions on the desired
relationship between The Canadian Field-Naturalist
and The Ottawa Field-Naturalists’ Club have led to
the recommendation that an ad hoc committee of
Council be set up to study the roles of our publications
and their relationship to the Club: and to produce a
discussion paper on journal policies and editorial
guidelines, to be brought before Council. As a result
of a desire to strengthen the relationship between
Council and the membership, a Council reporter was
appointed, whose articles on Council activities now
appear regularly in Trail & Landscape.
Compiled from committee reports
and Council minutes by
D. R. LAUBITZ, Recording Secretary
Auditor’s Report
To: Members of The Ottawa Field-Naturalists’ Club
We have examined the balance sheet of The Ottawa Field-Naturalists’ Club as at September 30, 1978 and
related statements of Profit and Loss. Our examination included a general review of the accounting procedures
and such tests of the records and supporting vouchers as were considered necessary in the circumstances.
In our opinion these financial statements present fairly the financial position of the organization as at
September 30, 1978 and the result of its operations for the period in accordance with generally accepted
accounting principles.
December 2, 1978
(Signed) F. M. BRIGHAM
G.J. WASTENEYS
1979 THE OTTAWA FIELD-NATURALISTS’ CLUB pi}
The Ottawa Field-Naturalists’ Club Balance Sheet
as at September 30, 1978
Assets
Current
Gashmancdhvermude posits she sacks gies we ceases $ 38,240.19
ACCOUMS TECOVADIG hea ook dbaoode ood net owed b ocoroo Ol 2 SDI
ACCTUGG WASTES sey eo cree ee Ae eo ee Nach ree Greer an 10,822.26
Rrepaldeexpenses;—_centemmMialll nya... ee oot eis S32 2242339 Sie
Investments
Canada Savings Bonds due October 31, 1978 ........... 10,700.00
Fixed at cost
FSU OITA TO taper Mak ARN cess tae Weeae Guede dad Qlci ead aapntiees a) Sees 52950
Wesseaccummlatedsdepreciation ase a. c-oee eees oe e 435.12 94.38
Total assets 64752012
Liabilities and Surplus
Current liabilities
AGCOWRIS DEVAS eine cere eh en ee oe ener $ 1,506.54
IDetierral: Tine aeons ee alga ean teenie oe e ee ene 7,675.83 $ 9,182.37
Surplus
Balancer @ ctobe wall wlO Te tar ses eieceec te ave cyaneisow stele cies enc 43,934.07
AGGE MCOMS Over Qos 5 socewsssoaouo gubleo do ue
he @ttawa Ficld=Naturalists: @lub)..... 4... - $ 4,155.60
hier Canadian-hicldsNatunalist.s esc la. 7,480.08 11,635.68
DATES SSvicnlysas) WIUS soocoesduceesedasccdn so ooos 35,5095)
Total lalbiiGes Aine SWRI Se de coeuoceaeoonceoamdsoud: $ 64,752.12
(Signed) Geoffrey Wasteneys, Auditor
F. Montgomery Brigham, Auditor
B. C. Henson, Treasurer
228 THE CANADIAN FIELD-NATURALIST Vol. 93
The Ottawa Field-Naturalists’ Club
Statement of Income and Expenditure
for the year ended September 30, 1978
Income
Apportionment of membership fees
PACTinier call Wrens eae ete ect cee, azar heed Wn cd asa, vad ehh ee $ 6,642.72
[STIS a es ie aa en Oe Ces se oe cess Beye ee ae ars nce oh ens Aa 720.00 3.7). 362072
iinailiscleandscape:—— SUDSCHIPtIONS) rae ess sie ee ee - 212.00
= Back mUmMbens sence as cccrtakee clone 35.00 247.00
ING® NOE SNA BCG Ore oO ODIO CRE COED EER ARC eae 442.78
Salles — decals, GmMleMMS oocccocvcccccuccgoooosouCb OOO 70.90
8,123.40
ISSUES 85 Gece aetin esl outches OIERES eno > US ERTIES RL TENS, oe 2,051.99 $ 10,175.39
Expenditure
Trail & Landscape
1? {Lo} WAS RTA Speer So, oer a ete ao RNa ene ee a Ba Deh)
Gigculatrone ryan eer ocr niet emis ae tien hea 119.61
Edit peancdrothices cats marches c.g setulae ace ees 54.09
[mI@ NIXON ANE, 5G oliaie Suck aco RR Se Buns Pec choke cheno ee cere nr oe ___ 440.00 3,191.40
Committee activities — net
EXCursiOnseanG MleChunes mum ern cene et ie ees oe eee (146.50)
NECN DEns Mp rte mmencccnestae conte sieesea ee ea aon se sacasia ae aus Ql 22
Miaic ou G hulp th arta, bears cms eek tener nian ae ae Lee 136.62
G@OnSenvanvOnbete eeepc rae eared ote tet ci eepers RTA ee 175.83
Gente rian alee eae vate eh yc shes Uy ge tht a 27.00
Bind@liced ercwanct ake cattery keke ER art Ree ee 590.11
ORC hVGRS UVC ar cnr wectaecicgan ac eee a eo Ee 40.83 1,781.11
BaldiwimyScholarships, ssc kns cca ee eee ak ek oe scutes 250.00
SPEClARACKIVITIES urs ahs ete nce tcstiarche eRe tan ae oi nests a aictac’ 114.91
GOuMCIIREPENSES pry er cans oe roe eae Oe cc ehererc kote 120.00
OHHICERS UD PICS teres ae ey he oe ae mee cette Mieka Tone 276.37
FLO MOVANT eas ee nee eee ena eG iene ata aes otioaets 200.00
MS Celeane@ USire tts se dete eek Ach ah ee enue hae LI 86.00 1,047.28
6,019.79
Excess of income over expenditure ...................... $ 4,155.60
1979 THE OTTAWA FIELD-NATURALISTS’ CLUB
The Ottawa Field-Naturalists’ Club
229
Statement of Income and Expenditure of The Canadian Field-Naturalist
for the year ended September 30, 1978
Income
Apportionment of The Ottawa Field-Naturalists’ Club Membership Fees
PNGNINU gee eet tae re eee eee ac eh SNe MR ose aa vane ohe ls $ 4,428.48
LUG so. gigs oNERSRe ne Een eres Ne te PER cere 480.00
4,908.48
SUMOSCHMIUTOMS teensy ei ccaus ayes tune wks S-oreye eieeaiet © sisieye Seis) werets 14,737.94
ublicationi—sREphINtS vcpusee ee ae 4 fem s oe oe yt «sete 5,761.00
= lalates andutalbssettines quer aryacine sicily 2,025.50
SpE XGA YP ARES sols eieueysueroney=fed-lepsrepaiahsiavora svoie 7,897.00
SB ACK SMUIMIDEES ints cscr-cereveseicics hee sicnstel onererene 1,803.73
Grants — National Research Council of Canada ........ 2,900.00
— Canadian National Sportsmen’s Show ........ 125.00
OMe TION CTES ita. cece sertnele se crete se tee nisin wie ses elsiasies 2,214.11
See MAINS Chaput stectens cccie venole e's aleuctic vena lenase, estates 962.13
Expenditure
Peatitbo SAVIN Merete ep cree ce ree cuesin stsnosecl ina tet aye aapcysGeteforeleltoren ot =ne 23,079.80
RGD TANN USHA SP chic arith ayicye {nv yrensyrays lasaysay stray SeeKoer olel Ble lasevalio'ss Se Sse = 3,073.09
GincwlatirOnyseryeetia ates whee cei sualere whale Glew ote siahsusisyore GS se ws
diinmeg ade xpeNnSeSiseysews cise aveach che avetebeiemeuey oie eo as choi shes
OihicemaSSISCAMt i) sas seers a Sestabdetas a testes tava slisieieleos Sarena tens
FAOSUAG Cemeteries ceersecnere | cei a Re Sisie ste etetetlehd. Sieitie iene aVolekesape
CHMGS. SUYD/TNSS serene lS a Aue en tenn Chenin gies ete eae Mtoe
AOI Nec inl emcee alec suSt wet cus rcita estes le okt cone cue ees sci age foga7eh sce, ueMORs
IMDS celle OU Shara ce ctspots cae steyey acess sunt caeeeiarcne cos soe eaetsncueneia es
Excess of income over expenditure .................0.65,
$ 19,646.42
17,487.23
3,025.00
3,176.24
26,152.89
3,627.09
I TSe23
1,891.50
896.05
570.45
1,502.00
33.60
$ 43,334.89
35,854.81
$ 7,480.08
Instructions to Contributors
Content
The Canadian Field-Naturalist is a medium for the
publication of scientific papers by amateur and professional
naturalists or field-biologists reporting observations and
results of investigations in any field of natural history
provided that they are original, significant, and relevant to
Canada. All readers and other potential contributors are
invited to submit for consideration their manuscripts
meeting these criteria. As the journal has a flexible
publication policy, items not covered in the traditional
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Canadian Field-Naturalist
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i 7
a
TABLE OF CONTENTS (concluded)
News and Comment 201
Book Reviews
Zoology: Where to find birds in British Columbia — The moths of America north of Mexico,
including Greenland: Fascicle 22.2, Noctuoidea (in part): Lymantriidae — Hawks,
falcons and falconry — Fishes of the world — Nesting ecology of Canada Geese
in the Hudson Bay Lowlands of Ontario: evolution and population regulation —
Birds in boreal Canada — Lexique anglais-frangais, termes techniques a usage des
biologistes — An introduction to the aquatic insects of North America — Nichoirs
d’oiseaux — Birds in peril: a guide to the endangered birds of Canada and the
United States
No
S
eS)
Botany: Manual of the vascular plants of Wyoming — A provisional checklist of species DAT
for Flora North America (revised) — Garden spice and wild pot-herbs: an American
herbal — How to identify grasses and grasslike plants (sedges and rushes) — Flora
of Alberta: a checklist — Freshwater wetlands: ecological processes and management
potential
Environment: Boreal ecology — The environmental impact of outdoor recreation — L’inventaire ZAltS
du capital nature: méthode de classification et de cartographie écologique du territoire
(3e€me approximation) — Applications de la télédétection a l’étude de la biosphere —
Wildness is all around us: notes of an urban naturalist
Other: Field photography: beginning and advanced techniques 218
New Titles 219
The Ottawa Field-Naturalists’ Club D8
Mailing date of previous issue 5 February 1979
1979 Council — The Ottawa Field-Naturalists’ Club
President: Roger Taylor Elisabeth Beaubien Fran Goodspeed
i : " pat Charlie Beddoe Peter Hall
Vice-President: Courtney Gilliatt aml Rekha Fe Nine conic
Treasurer: Barry Henson Frank Bell Frank Pope
2 } : Bill Cody lan Sneddon
Recording Secretary: Diana Laubitz Janel Dicemean Ken Strang
Corresponding Secretary: Valerie Hume Ellaine Dickson Ken Taylor
Loney Dickson Ewen Todd
Past President: Roger Foxall Marc Forget
Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists’ Club,
Box 3264, Postal Station C, Ottawa, Canada KIY 4J5. For information on Club activities telephone (613) 722-3050.
THE CANADIAN FIELD-NATURALIST Volume 93, Number 2 1979
Editorial
Our responsibilities as field-naturalists and biologists LORRAINE C. SMITH
Articles
Natural fires as an index of paleoclimate J. TERASMAE and N. C. WEEKS
Virgin Douglas Fir forest on Saturna Island, British Columbia
THOMAS P. SULLIVAN
Colonial-nesting Herring Gulls and Common Terns in northeastern Saskatchewan
HARRY A. STELFOX and GREGG J. BREWSTER
Epiphytes on White Elm, U/mus americana, near Thunder Bay, Ontario
P. BARCLAY-ESTRUP and RICHARD A. SIMS
Habitat utilization and population densities of the amphibians of northeastern Alberta
WAYNE ROBERTS and VICTOR LEWIN
Caribou distribution and group composition associated with construction of
the Trans-Alaska pipeline
RAYMOND D. CAMERON, KENNETH R. WHITTEN, WALTER T. SMITH, and
DANIEL D. ROBY
Interesting vascular plants from southeastern Yukon Territory
GEORGE W. SCOTTER and WILLIAM J. CODY
Notes
Fall foods of Common Snipe on the Copper River Delta, Alaska
STANLEY E. SENNER and P. G. MICKELSON
Ivory Gull colonies in southeastern Ellesmere Island, arctic Canada
F. FRISCH and W. C. MORGAN
Wind-caused death of Great Cormorant MARTIN K. MCNICHOLL and GEOFFREY G. HOGAN
Habitat selection by wintering Snowy Owls (Nyctea scandiaca)
M. Ross LEIN and G. A. WEBBER
Two-lined Salamander, Eurycea bislineata, in Labrador FRANCIS R. COOK and JOHN PRESTON
First Canadian record of a Flathead Catfish E. J. CROSSMAN and J. H. LEACH
Earthworm cocoons as a drift component in a southern Ontario stream
DONALD P. SCHWERT and KENNETH W. DANCE
Flowering plant phenology at Sheep Mountain, southwest Yukon Territory MANFRED HOEFS
Eggshell thickness in American shorebirds before and since DDT
MICHAEL L. MORRISON and LLOYD F. KIFF
Response of wintering Moose to mechanical habitat rehabilitation in Alaska M. SIGMAN
New localities for the Northern Spring Salamander and the Four-toed Salamander in
southwestern Quebec DAVID M. GORDON
First record of the Long-tailed Shrew (Sorex dispar) in New Brunswick
GORDON L. KIRKLAND, JR., DAVID F. SCHMIDT. and CAROL J. KIRKLAND
Intraspecific food theft by the American Kestrel PETER M. FETTEROLF
First record of the Northern Brook Lamprey, /chthvomyzon fossor, in the Nelson
River drainage, Manitoba J. JYRKKANEN and D. G. WRIGHT
Blue Grouse brood hen ~ Black Bear confrontation M. G. SULLIVAN
113
116
126
132,
139
144
155
163
195
198
199
200
concluded on inside back cover
ISSN 0008-3550
The CANADIAN
FIELD-NATURALIST
Published b
y THE OTTAWA F
IELD-NATURA
LISTS’ CLUB, O
, Ottawa, Canada
The CANADIAN
FIELD-NATURALIST
ublished by THE Z o) Fe FURALISTS A wa ana
Pp E OTTAWA FIELD-NATURA CEL Ott d
B awa, C a
THE movers
SS ee
Published by The Ottawa Fjeld-Naturalists’ Club
Editor:
ARTHUR GIBSON,
mntomological Branch, Devs!
OLTAWA-
ent of A wricullure,
Associate Editors:
FIARLAN I. SMITH. w. H. PIARRINGTON, Pp. A. TAVERNER,
Anthropology: Entomology. Ornithology.
Mi. Q. MALTE, ph.D.. M. ¥. WILLIAM, PHD. dus RE- LAMBE, ¥.G.S..
Botany - Geology- Palaeontology’
por, JonNMACOUN, MA. arto KLoTZ, TAs: Cc, GoRDON Hewits, BD Sc.,
Conchology- Metgorology- Zoalogy-
ee
CONTENTS:
—_——
On Cheneosaurus tolmanensis, @ new genus and species of
trackodont dinosaur from. the Edmonton cretaceous of
Alberta, BY Lawrence 4 117
sR
By John M. Coop 123
« feeding habits
; = M. Sternberg 129
Concer ario Cra}
ON Noa ee 130
Bird Notes. By L. Mcl. Verrill Age
The padson-Merrill press, Limited. dedicat a Special Tssue
qssumD MARCH 9TH, 1 tye :
Entered at Ottawa Post Office Se See Ob mailer 2 lO raptors—especially the Pereg 5 E
: grine Falcon
Volume 90, Number 3
July-September 1976
Centennial Year
Volume 93, Number 3
July-September 1979
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
Patrons
Their Excellencies the Governor General and Mrs. Edward Schreyer
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural
heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse
information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving,
maintaining or restoring environments of high quality for living things.
The Members of Council are listed on the inside back cover.
The Canadian Field-Naturalist
The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas
expressed in this journal, however, are private and do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or
any other agency.
Editor: Lorraine C. Smith
Assistant to the Editor: Donald A. Smith Book Review Editor: J. Wilson Eedy
Associate Editors
@) DBind A. J. Erskine George H. La Roi
E. L. Bousfield Charles Jonkel David P. Scott
Francis R. Cook Charles J. Krebs Stephen M. Smith
W. O. Pruitt, Jr.
Copy Editor: Marilyn D. Dadswell Chairman, Publications Committee: J. K. Strang
Production Manager: Pauline A. Smith Business Manager: W. J. Cody
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Subscription rates for individuals are $10 per calendar year. Libraries and other institutions may subscribe at the rate
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Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field- Naturalists’ Club, 1879-
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Business Manager: Mr. W. J. Cody, Box 3264, Postal Station C, Ottawa, Ontario, Canada KIY 45
Book Review Editor: Dr. J. Wilson Eedy, R.R. 1, Moffat, Ontario LOP 1J0
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Corvallis, Oregon, USA 97731 (address valid until August 1979).
Address manuscripts on birds to the Associate Editor for Ornithology:
Dr. A. J. Erskine, Canadian Wildlife Service, Box 1590, Sackville, New Brunswick EQOA 3C0
All other material intended for publication should be addressed to the Editor:
Dr. Lorraine C. Smith, R. R. 3, Stittsville, Ontario, Canada KOA 3G0
Urgent telephone calls may be made to the Editor's office (613-996-5840), the office of the Assistant to the Editor (613-231-
4304), or their home on evenings and weekends (613-836-1460), or to the Business Manager's office (613-995-9461).
Cover: Since The Ottawa Field-Naturalists’ Club was founded in 1879, it has published a scientific journal. Two covers are
illustrated. Left, The Ottawa Naturalist, published from 1887 to 1919. Right, The Canadian Field-Naturalist,
published from 1919 to the present; the format with a photograph on the cover was introduced in 1970.
The Canadian Field-Naturalist
Volume 93, Number 3 July-September 1979
To a Bigot
For years you have been telling us about the supreme importance of your laboratory studies in biology and we
have been convinced that your research is making contributions to scientific knowledge. But we have also
listened, sometimes with considerable repressed anger (because we know it is useless trying to argue with you:
“Don’t confuse me with facts, my mind is made up!”) as you denigrate any studies dealing with natural history.
Unfortunately, despite our vehement, logical protests, you refuse to hear the other side of the story but remain
adamant, being dogmatically convinced only of the correctness of your own beliefs and opinions. You think
further that investigations on whole animals or plants are merely diversions from what you consider to be
worthwhile science. But then you are not alone because there are many others like you.
Why am I writing this editorial? Well, I hope to warn others about your bigotry, especially because you are often
in a position to influence or make decisions regarding the research and careers of field-biologists. Indeed, I have
often heard you dismiss the merits of scientists’ publications as of “little scientific value” because they were
concerned with ecology, life history, taxonomy, or distribution of animals and plants, and were published in
“just naturalists’ journals.” You downgrade the worth of field-biologists but are firmly and smugly convinced of
your own worth.
In your narrow-mindedness you automatically equate the total amount of a research grant with the scientific
worth of the researcher. We think you are wrong. Everyone recognizes that some types of research are very
costly but these studies are not necessarily, and often are not, the most valuable and significant. If a researcher
can carry out his studies with a grant of x dollars, why should he ask for 3x or 30x dollars other than to gain more
prestige in the eyes of people like yourself? Some projects do not require grants of thousands and thousands of
dollars for very expensive materials and equipment. Fortunately some research, and it is often excellent
research, can be done by dedicated naturalists and field-biologists with a minimum of funds or even none at all.
Logically, value judgments should be made on the work accomplished and not on the size of the grants.
If you look at history, you'll find that some outstanding scientists carried out first-rate science both in the
laboratory and field with a minimum of technical help and materials. “Oh yes,” you'll say, “but times have
changed.” Molecular biology, and studies of the physiology and biochemistry of small bits of living protoplasm
or ultrastructures are the “in” fields but for these research projects (like yours) is good funding absolutely
essential, even at the expense of other biological research? How often have persuasive and persistent people like
you convinced granting bodies and governments to fund your research at enormous expense and cared little for
those who ask for a small share. In fact, when you think certain mission-oriented or applied problems are more
acceptable, haven’t you slanted your research grant proposal towards some aspect of cancer, energy, food
production, or the environment in order to increase your chances in the competition for available funds?
Some of us have been trained and have worked in your discipline as well as in field biology and consequently
we have an understanding of both these branches of biology. You, however, know only one. Although we doubt
you'll ever listen, you should be aware that knowledgeable and broad-minded biologists recognize the
importance of all good biological research. Of course we understand that you value your studies highly but we
cannot, and will not, accept your narrow-minded view that your researches are inherently a cut above others,
especially natural history and field studies. There is no doubt in our minds that investigations of plants, animals,
and whole communities are of the utmost importance. The more we learn about ecosystems and the more we
discover how plants and animals have adapted to their changing environments, the better we will be able to
conserve and manage our renewable natural resources, and to make predictions for the future. On the other
hand, as you learn in your research more and more about less and less, how can you even begin to understand
most of the natural world?
No doubt you won’t read this editorial because The Canadian Field-Naturalist to you is “just another
naturalists’ journal,” not a laboratory-oriented one and in your eyes doesn’t rate. But I hope other field-
biologists and naturalists will add their voices to mine so that in the end we will be able to counteract and destroy
the false premises on which your bigotry is built.
LORRAINE C. SMITH
Editor
231
Seasonal Growth, Food, and Feeding Habits of Young-of-the-year
Black Crappie in the Ottawa River
JOHN MARK HANSON and S. U. QADRI
Department of Biology, University of Ottawa, Ottawa, Ontario KIN 6N5
Hanson, John Mark and S. U. Qadri. 1979. Seasonal growth, food, and feeding habits of young-of-the-year Black Crappie
in the Ottawa River. Canadian Field-Naturalist 93(3): 232-238.
In the Ottawa River, during the first growing season, young-of-the-year Black Crappies (Pomoxis nigromaculatus) attained
an average total length of 68.4 mmanda mean weight of 4.03 g. The length-weight relationship was log WT = -4.84 + 2.97 log
TL. Young-of-the-year Black Crappies fed mostly on copepods and cladocerans from June to September but switched to
amphipods in October. They are diurnal in habit and feed mostly on organisms in the mid-waters of the littoral zone.
Key Words: growth, food organisms, feeding behaviour, Black Crappie, Pomoxis nigromaculatus, juveniles, copepods,
amphipods, seasonal variations, Gatineau, Quebec, Ottawa River.
Black Crappies, Pomoxis nigromaculatus, in the
Ottawa River are at the northern limit for the species
(Scott and Crossman 1973); the fish are restricted to
downstream of the Chaudiére Falls (McAllister and
Coad 1974). Throughout its range in the United States
little information is available on growth and feeding
of young-of-the-year. In Canada, the feeding biology
of the adult fish has been studied only by Keast and
Webb (1966) and Keast (1968).
The purpose of the present study was to investigate
the seasonal growth, food, and feeding habits of
young-of-the-year Black Crappies in the Ottawa
River.
30
20
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June July
Methods and Materials
The study site was a section of the Ottawa River,
Kettle Island Bay, a long narrow bay, 5 ha in area
(75°39’0’N, 45°28’8”W). The average depth of the
area studied was 3 m and the maximum was 6 m. The
bay is unique in that the sand is overlain with silt and
decaying detritus. The river current near the bay
mouth has an average velocity of 0.63 m/s (R. G.
Warnock, University of Ottawa, unpublished data)
but the current inside the bay depends on wind
direction and velocity. The water temperature at the
surface for May to October is given in Figure |. Water
hardness was 25-50 mg/L, oxygen 6-9 mg/L, carbon
Aug. Sept.
FiGuRE 1. Water temperatures for Kettle Island Bay in 1977.
M3y)
1979
dioxide 8-15 mg/L, and pH 7.1 + 0.3; suspended
solids vary greatly.
In the bay, emergent vegetation consists of Sagit-
taria latifolia, Scirpus sp., and Polygonum sp.; dense
stands of Elodea sp. and Ceratophyllum sp. with
scattered Potamogeton spp. occur from 0.2 to about
1.5m; Potamogeton spp. and Nymphaea sp., with
Elodea sp. and Ceratophyllum sp. growing under-
neath, occur on the two humps (at 1.5 and 2.0 m) and
from the sandbar to the south shore on the eastern end
of the bay. This sandbar is flanked by thick beds of
Vallisneria sp.
Sampling for Black Crappies was done by towing a
green 6.35-mm mesh seine, 1.83 by 30.5 m. One end
was fixed on shore as the other was towed out from
shore and back by a boat with a 20-HP outboard
motor. From the 2000 or more young captured on
each sample date, only 30-35 were kept; these were
sampled for lengths (mm), weight (g), scales, and
stomach contents. Sampling was done at 2-wk
intervals from May to October of 1977 and about 300
young were randomly selected for detailed examina-
tion. Organisms from the stomachs of fish captured in
the field were identified (mostly at order level) and the
number of organisms per fish, percent frequency, and
percent volume were compiled on a monthly basis.
Total length (mm) was used throughout. For com-
parison to other studies, conversion factors for fork
length and standard length were obtained as follows:
HANSON AND QADRI: YOUNG-OF-THE-YEAR BLACK CRAPPIE
Fork length (cm)
psy
NS
233
SL = 0.8302 TL — 0.3206 (based on 71 individual
measurements) and FL = 0.94 TL + 0.0135 (Figure 2).
Wet weights were taken to the nearest 0.01 g. The
anterior scale radius was measured with an ocular
micrometer (40 X) and reduced to actual size in
millimetres. Fulton’s condition factor was calculated
for each fish. The significance of the difference of
sample means was determined by a /-test.
Food and feeding habits of young Black Crappies
were observed in the field and in the aquarium. From
late June to early August, 10 young-of-the-year Black
Crappies were maintained in an aerated all-glass
aquarium, 61 X 30.5 X 30.5 cm. City of Ottawa water
(pH 6.9-7.2,; hardness 40-50 mg/L) was kept at
22+1°C. Substrate from Kettle Island Bay was
planted with Vallisneria sp., Potamogeton sp., and
Ceratophyllum sp. Lighting was from fluorescent
lights and sunlight.
In order to determine what foods could be eaten,
food preferences, and feeding chronology, the fish in
the aquarium were fed the following: Brine Shrimp
(Artemia salinas) nauplii, cladocerans, copepods,
amphipods (Hyallela azteca), free-swimming nema-
todes, dipteran (Chironomus sp.) larvae, white worms
(Enchytraeus albidus), vestigial winged fruit flies
(Drosophila melanogaster), newborn Guppies ( Poeci-
lia reticulatus), and shiner (Notemigonus crysoleucas
and Notropis spp.) fry. The young crappies were
observed for | h after the food was introduced.
| 2 3 4
5 6 7 8 9
Total length (cm)
FIGURE 2. Relationships of fork length to total length of young-of-the-year Black Crappies in 1977. FL = 0.9443 TL + 0.0135.
Averages of lengths with 95% confidence limits are shown.
234
Results
Large numbers of young-of-the-year Black Crappie
were captured near the Elodea and Ceratophyllum
beds until October when the weeds disintegrated and
the young moved into deeper waters. During the study
period all sizes of Yellow Perch (Perca flavescens),
Northern Pike (Esox Jucius), Largemouth Bass
(Micropterus salmoides), Pumpkinseed (Lepomis
gibbosus), Rockbass (Ambloplites rupestris), Brown
Bullhead (Jctalurus nebulosus), Golden Shiner ( Note-
migonus crysoleucas), Emerald Shiner (Notropis
atherinoides), and Silvery Minnow (Hybognathus
nuchalis) were captured consistently, but in varying
proportions, with the Black Crappies. The first batch
of young Black Crappie was captured 20 June 1977.
The mean length of the first sample of young Black
Crappie was 25.5 mm and rapid growth occurred until
mid-August when growth in length slowed to a value
of 68.4 mm (Figure 3). The length-weight relationship
(Figure 4) is exponential with a slope of 2.98.
Rapid growth in weight occurred from July to late
September (Figure 5) and slowed to a mean value of
4.03 g. The mean condition factor was 1.30 (Figure 6)
except for early August when the value of 1.01 is
Total length (cm)
Ts) S50) 15 3|
June July
THE CANADIAN FIELD-NATURALIST
Vol. 93
significantly (P< 0.005) lower than adjacent values.
Scales appeared on fish at about 25 mm in length,
weighing an average of 0.23 g. The relationship
between length and the anterior scale radius (Figure 7)
is given by Scale Radius = 0.1094 TL —-0.1023, show-
ing direct proportionality for young-of-the-year Black
Crappies.
Table 1 summarizes, by month, the food items
taken by young Black Crappies in Kettle Island Bay.
The percentage of empty stomachs was always less
than 9 and the stomachs containing food were usually
at least half full. Copepods occupied 48 to 90% of
stomach volumes from June to September but were
only 13% of the volume in October. Cladocerans
occurred to a maximum of 50% of stomach volume in
September but only 4% by volume in August.
Amphipods occurred maximally at 57% of the volume
in October.
Under conditions of high winds and/or roiled
water, young Black Crappies could not be observed
feeding. On very bright days, very few young Black
Crappies were observed feeding within 30 cm of the
surface, except in direct shade. Otherwise there was no
observed preference for position in the water column.
Sept.
FIGURE 3. The increase in length of young-of-the-year Black Crappies during the summer and fall of 1977. Averages and 95%
confidence limits are presented. Curve fitted by eye.
1979
LOgjo weight
0.1 02 03 04 05 06 07 08 09 10
LOgio total length
FIGURE 4. Length-weight relationship of young-of-the-year
Black Crappies in 1977. log WT = -4.84 + 2.965 log
TL. Length in millimetres, weight in grams.
During the feeding studies in the aquarium it was
noted that after sunset the young crappies remained
motionless near the bottom either in a corner or
among the vegetation. Only nematodes were entirely
ignored, although dipterans that reached the bottom
and amphipods that clung to weeds usually escaped.
Brine Shrimp nauplii were most readily consumed.
The agility of newborn Guppies made them
difficult to capture. All other foods offered were eaten
within the hour of observation.
Discussion
The mean total length of 68 mm for the first
growing season for Black Crappie in the Ottawa River
is comparable to values of 61 to 69 mm in Minnesota
(J. H. Keuhn, unpublished data; Moyle and Burrows
1954) and 68 mm in South Dakota (J. T. Shields,
HANSON AND QADRI: YOUNG-OF-THE-YEAR BLACK CRAPPIE
235
Weight (g)
June July Aug.
Sept. Oct.
FIGURE 5. The increase in weight of young-of-the-year Black
Crappies during the summer and fall of 1977.
Averages and 95% confidence limits are presented.
Curve fitted by eye.
unpublished data; Vanderpuye and Carlander 1971)
but are far smaller than those fish in southern ponds
(130-200 mm, Tucker 1973) and lakes (47-91 mm,
Erikson 1952). The northern growing season is shorter
and food is less abundant than in the more eutrophic
southern waters.
The length-weight relationship has a slope of
almost 3.0 and it does not vary significantly from this
value throughout its range (Table 2). The fairly
consistent condition factor (Figure 6) supports the
argument that body proportions of the young
crappies were constant during the study period. The
significant decrease in condition in early August
occurred after a 2'4-wk period of unusually cold, wet,
windy weather during which the young Black Crap-
pies were inactive. Contrary to our findings, Neal
(1963) found that temperature and water levels did not
affect the growth of Black Crappies in Clear Lake,
Iowa. The shorter growing season in the Ottawa River
would tend to exaggerate any condition affecting
growth. Table 3 shows that young-of-the-year Black
Crappies have essentially the same condition factor
throughout the range.
236 THE CANADIAN FIELD-NATURALIST Vol. 93
1.4 he $030
(24)
1.3 (35)
i ‘6 ~b01 fe, ten
(i)
Condition factor
[So eso lom esl 157 rs 15a SO anova
June July Aug. Sept. Oct.
FIGURE 6. Condition factor of young-of-the-year Black Crappies during the summer and fall of 1977. Averages and 95%
confidence limits are presented. Sample size is in parentheses.
TABLE |—Percent volume (above), percent occurrence (center), and number of prey items per fish (below) of the stomach
contents of young-of-the-year Black Crappie in 1977
July
August 9.86
September 9.86
6.97
October
1979
Anterior scale radius (mm)
Total length (cm)
FIGURE 7. The relationship of total length to the anterior
scale radius for young-of-the-year Black Crappies in
1977. Scale radius = 0.1094 TL -0.1203. Averages and
95% confidence limits are shown.
HANSON AND QADRI: YOUNG-OF-THE-YEAR BLACK CRAPPIE M37)
The body length to scale radius relationship was
directly proportional as was found for young crappies
in South Dakota (Vanderpuye and Carlander 1971)
and in Iowa (Erikson 1952). Later year classes may or
may not have direct proportionality (Carlander 1977).
The food eaten by young Black Crappies does not
seem to vary much throughout its distribution. Pearse
(1919) and Ewers (1934) reported that young Black
Crappies ate copepods and cladocerans in early
stages, with a progression to insects. Amphipods were
not an important food item in either of these studies
but the number of fish examined was small and some
of the fish may have been yearlings. The destruction of
the weedbeds, notably Elodea sp., in October seems to
account for the increased availability of amphipods as
a food item. This was also reflected in the increased
numbers of odonatans taken at the same time. The
relative unimportance of dipterans in the diet can be
attributed to their benthic habits and mostly noctur-
nal movements and the diurnal feeding habits of
young Black Crappies. Mathur and Robbins (1971)
found that young White Crappies (Pomoxis an-
nularis) are diurnal and eat much the same food items
as young Black Crappies. Although Reid (1949)
reported that Black Crappies in Florida do not eat fish
until they are about 80 mm long, Burris (1956) found a
25-mm individual that had eaten a fish fry. In this
study, a small percentage of Black Crappies from 25 to
TABLE 2—Length-weight relationships of young-of-the-year Black Crappies from various sources
Length-weight relationship
Source
Location Total length (mm)
Alabama Y oung-of-the-year
Oklahoma 15-56
South Dakota
South Dakota
Y oung-of-the-year
Y oung-of-the-year
Ottawa River 25-68
log WT = -4.710 + 2.914 log TL
log WT = -5.659 + 3.351 log TL
log WT = -5.252 + 3.198 log TL
log WT = -5.019 + 3.075 log TL
log WT = -4.835 + 2.97 log TL
Tucker (1973)
Burris (1956)
Nelson (1974)
Vanderpuye & Carlander
(1971)
Present study
TABLE 3—Condition factor of young-of-the-year Black Crappies from various sources
Location Total length (mm)
Alabama 50-155
Alabama Y oung-of-the-year
Oklahoma 15-56 (hatchery fish)
Minnesota (standard) —
Ottawa River 25-68
Condition factor Source
1.21 Swingle (1965)
1.00-1.45 Tucker (1973)
0.79 Burris (1956)
< 1.05 (poor)
1.22-1.50 (average)
> 1.88 (excellent) Carlander (1944)*
1.30 Present study
*Unpublished data with Minnesota Bureau of Fisheries Resource Investigations Report 41, revised, 40 pp., typewritten (quoted in Carlander
1977).
238
33 mm long ate fry. Both shiner fry and newborn
Guppies were eaten in the aquarium but the agility of
the Guppies suggests that shiner fry, too, may soon
become too difficult for young Black Crappies to
capture under natural conditions.
Literature Cited
Burris, W. E. 1956. Studies of the age, growth, and food of
known age young-of-the-year black crappies and of
stunted and fast growing black and white crappies of some
Oklahoma lakes. Ph.D. thesis, Oklahoma A & M College,
Miami. 88 pp. (quoted by Carlander 1977).
Carlander, K. D. 1977. Handbook of freshwater fishery
biology. Volume 2. Iowa State University Press, Ames,
Iowa. 431 pp.
Erikson, J. G. 1952. Age and growth of the black and white
crappies, Pomoxis nigromaculatus (Le Sueur) and Pomo-
xis annularis Rafinesque, in Clear Lake, Iowa. Iowa State
Journal of Science 26: 491-505.
Ewers, L. A. 1934. Summary report of crustaceans used as
food by the fishes of the western end of Lake Erie.
Transactions of the American Fisheries Society 64:
379-390.
Keast, A. 1968. Feeding biology of black crappie, Pomox-
is nigromaculatus. Journal of the Fisheries Research
Board of Canada 25: 285-297.
Keast, A. and D. Webb. 1966. Mouth and body form
relative to feeding ecology in the fish fauna of a small lake,
Lake Opinicon, Ontario. Journal of the Fisheries Re-
search Board of Canada 23: 1848-1867.
Mathur, D. and T. W. Robbins. 1971. Food habits and
feeding chronology of young white crappie, Pomoxis
annularis Rafinesque, in Conewingo Reservoir. Transac-
tions of the American Fisheries Society 100: 307-311.
McAllister, D.E. and B.W. Coad. 1974. Fishes of
Canada’s National Capital Region. Fisheries and Marine
Service, Miscellaneous Special Publication 24. 200 pp.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Moyle, J. B. and C. R. Burrows. 1954. Manual of instruc-
tions for lake survey. Minnesota Bureau of Fisheries
Research Unit Special Publication 1. 70 pp. (quoted by
Carlander 1977).
Neal, R. A. 1963. Black and white crappies in Clear lake,
1950-1961. Iowa State Journal of Science 37: 425-445.
Nelson, W. R. 1974. Age, growth, and maturity of thirteen
species of fish from Lake Oahe during the early years of
impoundment, 1963-1968. U.S. Fish and Wildlife Service
Technical Paper 77: 1-29. (quoted by Carlander 1977).
Pearse, A. S. 1919. Habits of the black crappie in inland
lakes of Wisconsin. U.S. Bureau of Fisheries Document
867: 5-16.
Reid, G. R. 1949. Food of the black crappie, Pomoxis
nigromaculatus, in Orange Lake, Florida. Transactions of
the American Fisheries Society 79: 145-154.
Scott, W. W. and E. J. Crossman. 1973. Freshwater fishes
of Canada. Fisheries Research Board of Canada, Bulletin
184. 966 pp.
Swingle, W. E. 1965. Length-weight relationships of Ala-
bama fishes. Auburn University Agricultural Experi-
mental Zoology Series in Fisheries 3. 87 pp. (qouted by
Carlander 1977).
Tucker, W.H. 1973. Food habits, growth, and length-
weight relationship of young-of-the-year black crappie
and largemouth bass in ponds. Jn Proceedings of the 26th
Annual Conference of South Eastern Game and Fish
Commissioners. Edited by A. L. Mitchell. pp. 565-577.
Vanderpuye, C.J. and K.D. Carlander. 1971. Age,
growth, and condition of black crappie, Pomoxis nigro-
maculatus (Le Sueur), in Lewis and Clark Lake, South
Dakota, 1954-1967. Iowa State Journal of Science 45:
541-555.
Received 25 November 1978
Accepted 22 January 1979
Nesting and Food-Storage Behavior of Peromyscus
maniculatus gracilis and P. leucopus noveboracensis
C. CRAIG TADLOCK and HAROLD G. KLEIN
Department of Biological Sciences, State University of New York, Plattsburgh, New York
12901
Tadlock, C. Craig and Harold G. Klein. 1979. Nesting and food-storage behavior of Peromyscus maniculatus gracilis and
P. leucopus noveboracensis. Canadian Field-Naturalist 93(3): 239-242.
In a laboratory study we compared the nesting and food-storage behavior of Peromyscus maniculatus gracilis and P. leucopus
noveboracensis (Rodentia: Muridae) given a choice between upper and lower nest-boxes. Peromyscus |. noveboracensis
preferred upper nesting locations while P. m. gracilis showed no significant preference between upper and lower nest-boxes,
although more lower nest-boxes were used. These results agree with those of a previous field study on the ecological
distribution of these subspecies where both inhabited an upland forest habitat providing both tree- and ground-holes.
Peromyscus |. noveboracensis stored significantly more food and tended to build heavier nests than P. m. gracilis. This may be
related to the higher preferred ambient temperature for P. I. noveboracensis and its more southern geographic distribution as
compared to P. m. gracilis. Where these subspecies occur in the same habitat, competition between them may be minimized
because of differences in height-preference for nesting and food-storage.
Key Words: Peromyscus, nesting, food-storage, height-preference, interspecific competition, sympatry, behavior, ecological,
geographical, physiological.
Previous studies have shown that the Deer Mouse,
Peromyscus maniculatus gracilis, and the White-
footed Mouse, P. leucopus noveboracensis, are
occasionally found living within the same woodland
habitats of northeastern United States and south-
eastern Canada (Klein 1960; Smith and Speller 1970).
Smith and Speller (1970) found both taxa cohabiting
an upland forest habitat in southeastern Ontario while
an adjacent mixed forest and a cedar forest were
occupied only by P. m. gracilis. They observed that a
significantly greater number of P. /. noveboracensis
ran up trees when released from traps, compared to
P. m. gracilis, which generally ran to ground refuges.
Because there were probably more tree-holes in the
upland forest than in the other forest habitats, one
hypothesis presented by these authors was that P. /.
noveboracensis occurred in the upland forest because
it could find refuge in the tree-holes present. The lack
of tree-holes in the mixed and cedar forests prevented
inhabitation by P./. noveboracensis because of
competition by P. m. gracilis for ground-holes.
The objective of this study was to determine
whether a difference in nest-height preference occurs
between these subspecies under laboratory condi-
tions. Differences in food-storing and nest-building
behavior, which might contribute to ecological sep-
aration of these taxa, were also investigated.
Material and Methods
Seven adult male Peromyscus m. gracilis and 10
adult male P./. noveboracensis were live-trapped
within 40 km of Plattsburgh, Clinton County, New
York, and were individually housed in plastic cages”
(25 X 10 X 15 cm) for 2-5 mo prior to testing. The
animals were housed and tested in a windowless, air-
conditioned room maintained at 20.6 + 2°C and with
a 13 light:11 dark cycle. A 7.5-W light provided dim
illumination during the dark phase. Food pellets
(Agway Charles River Rat and Mouse Formula) and
water were provided ad libitum.
The test apparatus was a modified version of one
used by Klein and Layne (1978). It consisted of a
pressed-board (Masonite) box 60 X 60 X 122 cm.
Eight removable nest-boxes (16 X 14 X 14cm) with
removable tops were attached to the outside of the
box, with four nest-boxes at each of two levels.
Circular entrance holes (5-cm diameter) to the upper
and lower nest-boxes were located 98 and 4 cm,
respectively, from the floor of the apparatus. Access’
to each upper nest-box was provided by a transversely-
grooved wooden stick which extended obliquely
within the apparatus from its floor to the nest-box
entrance hole. The apparatus had a removable
pressed-board floor and a wire mesh (7.5 mm) cover.
The floor of the apparatus was covered witha 7.5-mm
layer of a clay-based cat-box absorbent litter (Poise).
A food-dish and water-bottle were centrally located.
Two types of nesting material were provided. Pieces of
paper-towel material (13 X 2 cm) and pieces of non-
surgical absorbent cotton (2 + 0.5 g) were scattered
on the apparatus floor at the beginning of each test
period.
Each mouse was tested for a 7-night period. The 10
P. |. noveboracensis were tested consecutively prior to
testing the seven P. m. gracilis. Between tests of the
two subspecies, the entire apparatus was washed with
U9)
240
a mild ammonia solution and allowed to air-dry fora
period of 5 wk. Between tests of individual mice, the
apparatus was cleaned with warm water and the
locations of the nest-boxes were randomized. We
believe that these procedures eliminated any attrac-
tion or avoidance responses by the test animals to
specific nest-boxes or nest-box levels. The results were
inspected for any evidence of selection of previously
used nest-boxes by the test animals; none was found.
On Day |, at approximately 6 h into the light phase
of the daily cycle, each animal was put into the
apparatus through a lower-level entrance hole. On
Days 2 through 7, at about 6 h into the light phase, the
following data were recorded: location of mouse,
location of nest(s), location and number of food
pellets, and location and number of feces. On Day 8,
at 4h into the light phase, the animal was removed
from the apparatus. In addition to the above data, the
weights of the nests and stored food were recorded.
The apparatus was cleaned and a new mouse was
introduced.
Results
Every mouse visited each of the eight nest-boxes, as
determined by the presence of feces in the nest-boxes.
Maximum-minimum thermometer data indicated no
difference in temperature between upper and lower
nesting levels, eliminating this factor as a possible
explanation for the nesting patterns observed in this
study.
Peromyscus |. noveboracensis exhibited a clearcut
preference for upper nest-boxes but P. m. gracilis
individuals were divided in their choice of nest-box
level (Table 1). Individually, the mice of both taxa
were highly consistent in their nestbox-level selection
during the 7-night period. None nested at both levels
of the apparatus and each mouse could be classified as
either an “upper” or “lower” nester (Table 1).
Nest construction was similar in both subspecies.
All nests were constructed of shredded cotton; no
THE CANADIAN FIELD-NATURALIST
Vol. 93
paper-towel material was incorporated into any of the
nests. The nests were all built in a spherical form, with
a small cavity in the center and with two openings.
A comparison of the nest weights of the two
subspecies, regardless of level, indicated a tendency
for P. 1. noveboracensis nests to weigh more than
P. m. gracilis nests (Table 1), but the difference was
not significant (Mann-Whitney U test, P= 0.094).
This was caused partly by the great variability in nest
weights, which ranged from 17.9 to 76.2 g in P. 1.
noveboracensis and from 9.5 to 53.2 g in P. m.
gracilis. A comparison of the nests built in the upper
boxes showed that P. /. noveboracensis nests were
significantly heavier than P. m. gracilis nests (Table
1). No significant difference was found between
weights of upper and lower nests of P. m. gracilis
(Mann-Whitney U test, P= 0.228).
Eight of the 10 P. 1. noveboracensis constructed
only one nest during the study period and these nests
were used in calculating the mean nest weight (Table
1) in order to make it comparable to the mean nest
weight for P. m. gracilis, which was calculated from
seven single nests. The other P. /. noveboracensis
individuals built two and three nests each within the 7-
night period, respectively. Each of these nests weighed
less than the mean weight of the eight single nests, and
their weights were not included in the calculation of
the mean nest weight because of this bias.
Food stores were usually independent of nests.
Only one P. m. gracilis stored food in its nesting-box.
Five P. /. noveboracensis stored food in their nesting-
boxes; of these, four individuals had additional food
stores, from three to six each. Peromyscus |. nove-
boracensis stored significantly more food than P. m.
gracilis (Table 2). Intrasubspecific comparisons of
weights of food stored in upper versus lower boxes
showed no significant differences in either subspecies
(Table 2). Although there was a tendency for P. /.
noveboracensis to store more food in upper than in
lower boxes, and for P. m. gracilis to store more food
TABLE |—Summary of nesting data for mice of each subspecies nesting at each nest-box level during the 7-night study period
ee SS ee a ee Se ee ere a IE
Nest
Subspecies location
P. m. gracilis Upper
Lower
P. |. noveboracensist Upper
Lower
Weight of all
Number of nests for each
mice Nest weight (g) subspecies (g)
Spee Mean + SE Mean + SE
4 IBS Se DSP
+
3 34.0 + 12.29 mae
2b *
. SOO ree 35.6 + 6.7
oooooooooo—owooaovwoaoasw@@w9w9wTw0DaO=oOD®ooo SESE eo
*Difference between these weights was significant at P = 0.016; Mann-Whitney U test (Siegel 1956).
tOnly eight single nests were used for calculating mean nest weights. See text for explanation.
1979
TADLOCK AND KLEIN: PEROMYSCUS NESTING BEHAVIOR
241
TABLE 2—Summary of food-storage data for mice of each subspecies storing food at each nest-box level during the
7-night study period. N = 7 for P. m. gracilis; N = 10 for P. 1. noveboracensis
Weight of Weight of total
Mean food stored food stored
Food-store number of at each level (g) by subspecies (g)
Subspecies location food-stores Mean + SE Wien xe Se
i Upper 1.1 9.58 + 5.84** in A
P. m. gracilis ee 26 18.55 + 5.65 28.13 + 10.90
; ; Upper DD 45.46 + 13.94** me cs
P. 1. noveboracensis es 19 30.38 + 15.54 75.84 + 17.38
*Difference between these values was significant at P< 0.05; Mann-Whitney U test (Siegel 1956).
**Difference between these values was significant at P= 0.018; Mann-Whitney U test (Siegel 1956).
in lower boxes, these differences were not significant
(Table 2). A comparison of food-storage between the
subspecies at each level showed that in the upper
boxes P. /. noveboracensis stored significantly greater
weights of food than P. m. gracilis (Table 2). In the
lower boxes, the weights of food stored by P. /.
noveboracensis were not significantly higher than
those stored by P. m. gracilis.
Discussion
The three variables (nest-height preference, nest
weight, and amount of food stored) measured in the
artificial environment of our study indicated a
significantly greater usage of upper nest-boxes by
Peromyscus |. noveboracensis compared to P. m.
gracilis. With respect to nest-height preference, the
choice of upper nest locations by P. /. novebora-
censis was unquestionable; a preference for upper-
level food storage sites, although not significant, was
relatively strong. This agrees with Nicholson’s (1941)
finding that a southern Michigan population of P. /.
noveboracensis used tree nest-boxes (0.9 to 9.2m
above ground) to a much greater extent than ground-
level boxes.
Peromyscus m. gracilis showed no significant
preference for nest-box level in nesting or food
storage, although heavier nests were built and more
food was stored in lower boxes. The patterns of nest-
box utilization by these subspecies in our artificial
environment supports the hypothesis of Smith and
Speller (1970) that cohabitation of these subspecies in
their upland forest habitat was based, in part, on
differences in the utilization of above-ground cavities.
Implicit in Smith and Speller’s hypothesis was the
assumption that P. /. noveboracensis was forced to
occupy arboreal situations because of competition for
ground holes from P. m. gracilis; however, the con-
sistent preference of P. /. noveboracensis for upper
boxes for nesting in our study suggests that this may
be a behavior typical for this subspecies. Therefore,
the observed arboreal tendencies of P. 1. novebora-
censis in the Smith and Speller (1970) study may not
have been a change in the behavior of this population
because of competition from P. m. gracilis.
The lack of nest-height preference by P. m.
gracilis in our study suggests an ability of this taxon to
adjust to various nesting situations. It agrees with the
widespread distribution of P. m. gracilis in the Smith
and Speller (1970) study and the observation that
P.m. gracilis individuals ran up trees on some
occasions, although they usually retreated into
ground holes. It also suggests that P. m. gracilis would
not be a serious competitor of P. /. noveboracensis
for arboreal nest sites except in periods of high
population density in P. m. gracilis. The low
population density of P. m. gracilis in the Smith and
Speller (1970) study, coupled with the tendency of
released mice to use ground refuges, is in agreement
with this inference.
Although P. /. noveboracensis stored a significantly
greater amount of food than P. m. gracilis, and
P. 1. noveboracensis tended to build heavier nests than
P. m. gracilis, it is possible that these differences
(mainly attributable to the apparent inhibition of
P. m. gracilis in its utilization of upper nest-boxes)
were not entirely behaviorally-based. The thermo-
regulatory function of the nest in Peromyscus has
been well-documented (Sealander 1952; Glaser and
Lustick 1975). A difference in ambient temperature
preference has been experimentally demonstrated
between P. /. noveboracensis (32.4°C selected) and
P. m. gracilis (29.1°C selected) by Ogilvie and Stinson
(1966). The differences between these preferred
ambient temperatures and the ambient temperature of
our study (20.6°C) were probably sufficient to elicit
temperature-induced food-storing and nest-building
responses in these animals. A lowered ambient
temperature (7° versus 27°C) caused an increase in
242 THE CANADIAN FIELD-NATURALIST
food-storing behavior in P. /. noveboracensis (Barry
1976). An ambient temperature of 20°C elicited a
maximal paper-shredding, nest-building response in
P. |. noveboracensis, compared to the responses at 25°
and 30°C (Jaslove and McManus 1972). The dif-
ferences in food-storing and nest-building behavior
shown by the subspecies in our study may have been
differential responses to the low ambient temperature
of our study relative to the preferred ambient
temperatures of these taxa. The differences in pre-
ferred ambient temperature is presumably based ona
physiological difference between the subspecies. Riedel
(1967) demonstrated that, at low temperatures, P. m.
gracilis survived better than P. 1. noveboracensis,
whereas the opposite was true at high temperatures.
A situation of severe competition for nest sites may
exist, however, between P. Jeuwcopus and a closely
related species, P. gossypinus, where they are sym-
patric in the northern part of the Gulf coastal plain
and in the Mississippi Valley of southern United
States. McCarley (1963) concluded that in this area
P. gossypinus excluded P. leucopus from lowland
forest habitats by interspecific competition. Com-
petition for arboreal nest-sites may have been a major
factor in this interspecific relationship. Ten male
P. gossypinus palmarius from Florida showed a very
strong preference for upper nest-boxes when tested in
an apparatus similar to that of our study (Klein and
Layne 1978). A direct comparison (using a similar
apparatus) of the nest-height preferences and inter-
actions of individuals from sympatric populations of
P. leucopus and P. gossypinus might clarify the extent
and nature of their interspecific competition.
The behavioral and physiological differences be-
tween P. m. gracilis and P. |. noveboracensis are
probably based on adaptations related to the geo-
graphic-ecologic distributions of these subspecies
(and their respective species). For example, nesting in
an arboreal site would be more appropriate in areas of
mild winter climate, whereas in areas of severe winter
climate it might cause a serious thermoregulatory
problem. Such a species- or subspecies-typical be-
havior, if unmodifiable, could be a factor limiting the
geographic-ecologic distribution of the taxon. The
different nest-height preferences shown by these
subspecies may reflect the climatic conditions in their
respective environments. Although the ranges of P. /.
noveboracensis and P. m. gracilis broadly overlap,
P. |. noveboracensis is more southern than P. m.
gracilis in its geographic distribution and, where these
taxa are sympatric, corresponding ecologic dif-
ferences in local distribution generally occur (Hamil-
ton 1943; Klein 1960; Miller and Kellogg 1955).
Where cohabitation of these subspecies occurs, our
study suggests that sufficient differences exist in the
height-preference aspect of nesting and food-storage
Vol. 93
behavior to result in decreased competition between
them.
Acknowledgments
We thank James N. Layne for the valuable
suggestions he made during the preparation of this
paper.
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Received 2 November 1978
Accepted 8 February 1979
Life History Characteristics of Little Brown Bats
(Myotis lucifugus) in Alberta
DAVID B. SCHOWALTER,! JOHN R. GUNSON,! and LAWRENCE D. HARDER?
'Alberta Fish and Wildlife Division, 6909-116 St., Edmonton, Alberta T6H 4P2
2Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9
Schowalter, D. B., J. R. Gunson, and L. D. Harder. 1979. Life history characteristics of Little Brown Bats (Myotis
lucifugus) in Alberta. Canadian Field-Naturalist 93(3): 243-251.
Life history data of Little Brown Bats (Myotis lucifugus) were recorded from 1971 to 1978 during the course of a rabies study
in Alberta. The study is the northern-most examination of the life history of the species to date. The number of bats that
inhabited the 269 maternity colonies examined, varied from less than 15 to approximately 1100 adults. Seventy-five percent of
the colonies in central Alberta were in occupied buildings. Differences in the biology of Little Brown Bats from more southern
studies included later commencement of parturition, shortened period of parturition, non-breeding of juvenile females,
substantial numbers of females at non-maternity shelters during mid-summer, extensive roosting at exposed locations during
summer and fall, and aggregations of unknown function in abandoned buildings during fall. Adult females gained weight
during and after pregnancy and in late summer, and lost weight at parturition and in late July. Juvenile males entered
hibernation weighing less than adult males. Adult males were seldom located in maternity roosts but predominated at a
hibernaculum.
Key Words: Little Brown Bat, Myotis lucifugus, life history, Alberta.
The Little Brown Bat (Myotis lucifugus) is one of
the most widely distributed species of North Ameri-
can bats (Hall and Kelson 1959); its range encom-
passes a wide variety of habitats and climates.
Latitudinal gradients in photoperiod and the duration ©
of the seasons could induce selective pressures that
might result in behavioral and physiological variation
within this species. The biology of M. /. lucifugus in
eastern North America (reviewed by Humphrey and
Cope 1976) and of M. 7. occulatus in New Mexico
(O’Farrell and Studier 1973, 1975; Studier and
O’Farrell 1972; Studier et al. 1973) have been
extensively studied, but life history information for
more northern populations of Little Brown Bats is
necessary to identify and define characteristics that
vary with latitude.
The initiation of a research program on bat rabies
was prompted by an outbreak of rabies in Alberta in
1970 and the first diagnosis of this disease in a bat
from the province in the following year. Dorward et
al. (1977) reported preliminary conclusions regarding
the incidence of rabies in bats from Alberta. This
paper presents biological data on the Little Brown
Bats collected during the course of the study. Observa-
tions were made from 49°N to 57°N, but most
information was collected between 52°N and 55°N,
the region commonly known as ‘central Alberta.’ This
area is 4-6° north of studies in Ontario (Fenton 1969,
1970), approximately 11° north of studies in Indiana
and Kentucky (Humphrey and Cope 1976), and
approximately 14° north of studies in New Mexico
(O’Farrell and Studier 1973, 1975).
The Little Brown Bats that inhabit the non-
mountainous portions of Alberta have been identified
as M. 1. lucifugus (Hall and Kelson 1959; Soper 1964;
Banfield 1974); however, southern Alberta is occupied
by a separate form, M./. carissima (Smith and
Schowalter 1979). Unless otherwise indicated and
with the exception of a discussion of the provincial
distribution of M. lucifugus, the data presented here
deal with M./. lucifugus as it occurs in central
Alberta, although some observations may be from
areas of intergradation with the southern form. Also
discussed are bats from a hibernaculum near Cado-
min (see Figure 1) which have been previously
classified as M./. pernox (Soper 1964) but are
indistinguishable from M./. lucifugus of central’
Alberta (H. C. Smith, Provincial Museum of Alberta,
personal communication).
Methods
Most colonies and some shelters were located
through complaints of bats made to government
offices by the public. Systematic surveys, responses to
newspaper advertisements, and a questionnaire
mailed to rural landowners provided additional
locations. History sheets of bats sent to the Animal
Diseases Research Institute (Western) [ADRI(W)] of
Agriculture Canada at Lethbridge, Alberta for rabies
testing provided the majority of the locality records
for individual bats. Additional individual records
were obtained from specimens at the Provincial
Museum of Alberta, the Museum of the Department
of Zoology at the University of Alberta, as well as
243
244 THE CANADIAN FIELD-NATURALIST
Rocky
200 km
Individual O
Colonies —
estimated size
Solonetzic
Soils
1-25 e
(78)
26-100 ®©
(99)
101-250 @
(57)
251-500 @
(28)
over — 500
(5)
Vol. 93
Suntain HouseO
FicuRE 1. Distribution and estimated size of Little Brown Bat maternity colonies located in Alberta, 1971-1977.
from published data and our own field activities.
Most roosts were entered to identify and enumerate
the bats present. Bats were periodically collected at
maternity roosts for rabies testing and additional bats
were captured at maternity roosts known to harbor
rabies, a variety of non-maternity shelters, a hiber-
naculum, and by mist-netting over ponds and streams.
Aggregations of bats in buildings in fall (‘fall aggrega-
tions’) were sampled in 1975. The following informa-
tion was recorded for each bat: age (juvenile or adult)
as determined by the closure of the epiphyses of the
finger joints, sex, and weight. Examination of the
nipples of adult female bats (Racey 1974) provided an
indication of their reproductive history, and the
current reproductive condition of a sample of female
bats taken prior to the commencement of parturition
was determined by uterine examination. The number
of fetuses and the horn of implantation were recorded
for pregnant females. Eight non-parous unsuckled
female bats collected at shelters in mid-June were aged
by dental annuli (Schowalter et al. 1978b) to
determine the ages of non-breeding females.
1979
Results and Discussion
Distribution
We examined 269 maternity roosts of Little Brown
Bats. Sufficient data to determine the relative abun-
dance and distribution of this species are available
only from the more heavily settled southern and
central areas of the province. Concentrations of
colonies were evident in several lake-cottage develop-
ments, in certain rural areas of knob and kettle (‘pot-
hole’) topography in central Alberta, and in areas of
extensive irrigation development near Brooks and
Lethbridge. Few colonies were located near Vegre-
ville, where the habitat resembled that of areas with
many colonies, or north of Brooks and Medicine Hat
(Figure 1). Questionnaire responses, interviews with
residents, and the pattern of submission of rabies-
suspect bats all suggested that few Little Brown Bats
are found in these regions. Silver-haired Bats ( Lasion-
ycteris noctivagans) (Schowalter et al. 1978a) and
Hoary Bats (Lasiurus cinereus) (Schowalter and
Dorward 1978) have been submitted to ADRI(W)
from these areas with sufficient frequency to suggest
that the absence of Little Brown Bats is probably not
an artifact of the reporting system.
The area of low abundance of Little Brown Bats in
Alberta corresponds, in part, to the distribution of
solonetzic soils (Figure 1) and the distribution of
grasslands (see Hardy 1967). Water bodies in solon-
etzic soil zones are known to support different
arthropod faunas than those of other areas (Rawson
and Moore 1944) and the food supply for bats in
solonetzic soil areas may thus be of different quality
than in other areas. Grasslands are less heavily settled
and have far fewer trees than other agricultural areas
and thus offer fewer potential roost sites. These
factors do not, however, appear to explain completely
the observed distribution of Little Brown Bats.
Maternity Colonies
Although some complaints of bats in cottages were
not investigated and colonies in unoccupied buildings
and natural roosts were less likely to be reported,
occupied buildings were apparently favored for
colony formation in central Alberta. Of the 196 M. /.
lucifugus maternity colonies located and classified in
this region, 147 were in occupied buildings and, with
the exception of one colony in a tree, the remainder
were in cottages, abandoned houses, sheds, and barns.
Maternity roosts were typically dark and poorly
ventilated and almost always subject to heating by the
sun, as described by Fenton (1970) and Humphrey
and Cope (1976). Most colonies occupied a single
building, although bats in larger colonies appeared to
use as many as six buildings. The estimated number of
adult Little Brown Bats in individual maternity
colonies ranged from less than 15 to 1100; between 50
SCHOWALTER ET AL.: LITTLE BROWN BATS, ALBERTA
245
and 300 was usual.
From 1972 to 1975 bats returned to the maternity
roosts during the first half of May. During 1976 and
1977, years with early and mild springs, bats were
reported as early as 24 April (1976) and many roosts
contained bats by | May. The earliest date that a
juvenile was taken at a non-nursery roost was 9 July
(1976), but juveniles and suckled females were
regularly observed at such locations only during the
last week of July, indicating that nursery break-up
was advanced by that time. By the second week of
August most maternity roosts had few bats, but
juveniles were found at this type of roost as late as 26
August (1975). Juvenile bats outnumbered adult
females in our August samples from maternity
colonies (69:30), indicating that adults tend to leave
nurseries sooner than juveniles, as Humphrey and
Cope (1976) have reported.
The sexes were equally represented in our collec-
tion of fetuses and juveniles from maternity colonies,
but adult males were seldom found in these roosts
(Table 1). This low occurrence of males agrees with
the observation by Humphrey and Cope (1976) that
males rarely frequent maternity colonies in the
northern portion of the species range compared with
regions to the south.
TABLE |1—The percentage of males among samples of Little
Brown Bats from central Alberta: 1972-1978.
Sample size in parentheses
Percentage of males
Sample type Fetuses Juveniles Adults
Maternity roost 53 (61) 49 (395) 0.4 (807)
Shelters
May 3 (Sl)
June 41 (66)
July 20 ~=(5) 38@8))
Aug. 31 (45) 24 (49)
Sept. 41 (106) 30 (147)
Fall aggregation 38 = (8) Li CS)
Swarming 67 (30) 80 (81)
Hibernation 78 (270)
Other species of bats were seldom encountered in
Little Brown Bat maternity roosts. Three attics used
by Little Brown Bats were also colonized by Big
Brown Bats (Eptesicus fuscus). Little Brown Bats
appeared to have recently colonized two of these attics
after modifications to the buildings and the two
species used different parts of the attics. In the third
case a single beam separated the roosts of the species,
but at the time of observation the majority of the Big
Brown Bats were in the walls of the building rather
than immediately adjacent to the Little Brown Bats. A
246
Silver-haired Bat was observed in one colony on 17
May (1974).
Shelters
We recognized two types of non-nursery roosts or
shelters used by Little Brown Bats. Some bats
persistently used concealed locations in barns, attics,
log buildings, piles of lumber, and behind shutters,
tree bark, and a piece of tin wrapped around a tree
trunk. One banded male was observed on each of six
inspections of a roost over three summers. Shelters in
exposed situations, typically darkened recesses of
brick buildings between 2 and 7 m above the ground,
were used daily by | to 22 bats. There was evidently a
high turnover of individuals at these locations: some
property owners destroyed roosting bats daily,
seemingly without reducing the number of bats using
the roost. Bats banded at this type of shelter have not
been recaptured at the original banding site.
More bats were collected at exposed roosts at night
than during the day. Although bats roosting at
shelters during the night usually became torpid, many
of those that were not disturbed, aroused and
departed prior to sunrise. Bats found during the day
were almost always torpid.
With the exception of a male collected on 23 May
1978, all bats captured at shelters during May were
females (Table 1); many had previously suckled young
(Table 2) and may have been migrating to maternity
colonies. Male Little Brown Bats were also poorly
represented (1/24) in May submissions to ADRI(W)
from Alberta and May samples from shelters in
Michigan (Miller 1955). These observations probably
relate to the tendency of males to depart from the
hibernacula later than females (Humphrey and Cope
1976).
TABLE 2—The percentage of unsuckled adult female Little
Brown Bats among samples from central Alberta: 1972-
1978. Sample size in parentheses
Percentage
Sample type Period unsuckled
Maternity roosts Prior to 15 June! 17 (329)
After 1 July? 18 (331)
Shelters May 30 (30)
June 95 (39)
July 88 (8)
Aug. 41 (37)
Sept. 44 (91)
Fall aggregations Aug.—Sept. 18 (17)
‘Pregnant unsuckled bats taken after 15 June could have
been identified as suckled.
2Bats which appeared to be unsuckled after | July are not
considered likely to give birth that year.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Female Little Brown Bats were observed at shelters
in Alberta proportionately more frequently than has
been reported elsewhere (Table 3). The proportion of
females appears to increase with latitude (Table 3),
but the distance from hibernacula is known to affect
the proportion of males in maternity roosts (Humph-
rey and Cope 1976) and may affect the sex ratio in
shelters. Female Little Brown Bats that use shelters
during June have been thought to be non-parous
(Miller 1955; Humphrey and Cope 1976). In Alberta
the majority of females taken at shelters during June
were unsuckled (Table 2). None of eight females
captured at shelters, whose uteri were examined was
pregnant. Palpation of other female bats at shelters
during June produced no evidence of pregnancy.
Although sex-ratio data for bats caught at shelters
(Table 1) represent observations made at varying
intervals with varying effort over 5 yr, significant
annual variation is assumed not to have occurred.
TABLE 3—Percentage of males among Little Brown Bats
taken in shelters during June from various North American
locations
Latitude Percent
Location north males (n) Reference
Oklahoma 35 100 (17) Glass and Ward (cited in
Humphrey and Cope
1976)
Indiana 39 100 (3) Humphrey and Cope 1976
West
Virginia 39
New York 42
100 (40) Krutzsch 1961
87 (63) Davis and Hitchcock 1965
Michigan 42 84 (85) Miller 1955
Vermont 42 98 (648) Davis and Hitchcock 1965
Alberta 54 41 (66) This study
The use of shelters increased during late July with
the appearance of juvenile and suckled adult female
bats. The largest numbers of bats were collected at
shelters around the end of August and generally
declined thereafter. During August and September
bats frequented roosts not used earlier in the year.
Bats roosting on white buildings were easily seen from
roadways and were collected by driving through
business districts. Exposed roosting was observed or
reported from throughout central Alberta. Bats
roosting at such locations may have been migrants
passing through unfamiliar areas. An adult male
Little Brown Bat banded at such a shelter in
Edmonton on 8 September 1977 was recaptured 5 d
later near Rocky Mountain House, a movement of
173 km.
1979
Frequent use of exposed roosts by Little Brown
Bats has not been reported previously. The roosting is
so obvious that it is doubtful that it would be missed
by bat researchers if it occurred in their research areas.
Efforts to locate bats or bat droppings at potential
exposed shelters in Lethbridge and Medicine Hat did
not produce evidence of bats, suggesting that the
subspecies found in southern Alberta may not use
exposed roosts. The behavior and transience of bats at
shelters were similar to those of the Cave Bat (Myotis
velifer) at ‘transient colonies’ in early spring and late
autumn in Kansas (Kunz 1974). This behavior is
thought to be important in reducing metabolic
expenditure and in migration. In central Alberta we
have taken a few Keen’s Bats (Myotis keenii) and Big
Brown Bats at shelters used by Little Brown Bats, and
Silver-haired Bats are frequently encountered by
window washers in exposed situations on higher
buildings in Edmonton. As similar observations in
other areas are not known to us, exposed roosting in
central Alberta may be important in_ thermo-
regulation, possibly in response to unidentified
climatic factors.
Pregnancy
Suckled
‘a
E
©
b&
co)
=
£
=
C)
s
>
a]
°
a
SCHOWALTER ET AL.: LITTLE BROWN BATS, ALBERTA
247
Fall Aggregations
We have observed concentrations of 500 or more
Little Brown Bats in buildings on three occasions
during late August and early September. All of the
buildings used by these ‘fall aggregations’ were
abandoned and unheated, but the bats were not torpid
during the day. Local residents reported that one of
these buildings had been used by bats during several
autumns. The predominance of heavy female bats in
two small samples (Table 1, Figure 2) suggests that fall
aggregations may be related to migration rather than
to breeding. Humphrey and Cope (1976) noted
evidence of “autumn dispersal and transient nursery
visitation” distinct from swarming which takes place
at caves. Fall aggregations in Alberta may be part of
the same phenomenon noted by Humphrey and Cope
(1976), but few data are available on activity of Little
Brown Bats away from caves at this time of year.
Weight Dynamics
Adult female Little Brown Bats gained weight
during pregnancy, after parturition, and prior to
hibernation (Figure 2). Weight losses occurred as a
8
Fall
Aggregation
Lactation
MAY JUNE
JULY AUGU ST SEPTEMBER
FIGURE 2. Seasonal variation in body weight of adult female Little Brown Bats in central Alberta (mean + SE, range),
1971-1977.
248
result of parturition and from unknown causes at the
end of July, at least among females in maternity
roosts. Data from Ontario (Fenton 1970) are
comparable to those presented here, but they do not
extend into August, so that it is not known whether
the weight loss observed at that time is general for the
species.
Juvenile Little Brown Bats gain weight rapidly until
late July (Figure 3). Stomachs of juveniles examined
at that time contained insects only, insects and milk,
or milk only, indicating that weaning had begun,
Weights are less consistent after that time, suggesting
that juveniles may lose some weight after colony
break-up. A collection on 20 August (Figure 3)
included many juveniles that appeared to be starving
and were of low weights.
Swarming and Hibernation
A preponderance of males was netted at the mouth
of Cadomin Cave during September 1977 (Table 1), a
finding that is similar to observations elsewhere
(Davis and Hitchcock 1965; Hall and Brenner 1968;
NURSERY
9
8
:
9 7
>
io 6
=
~
8
AS 5
4 20
NON-NURSERY
JUNE JULY AUGUST SEPTEMBER
Month
FIGURE 3. Seasonal variation in body weight of juvenile
Little Brown Bats in central Alberta (mean + SE,
range), 1974-1977.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Fenton 1969; Humphrey and Cope 1976). Swarming
may have ended at Cadomin Cave between 20 and 27
September 1977 as only four individuals were
captured on the later date. This change was probably
not caused by weather as Long-legged Bats (Myotis
volans) were captured in greater numbers on 27 than
on 20 September.
Juvenile bats captured on 20 September 1977 at
Cadomin Cave weighed significantly less (t-test,
P<0.01) than juveniles at shelters in Edmonton and
Edson on 19 and 20 September (Table 4). Weights of
adult males in the two samples were nearly identical.
Too few adult females were captured at Cadomin
Cave to allow comparison. Juveniles may have lost
weight in moving to the cave from maternity roosts,
whereas the distance moved by adult males from the
summer range may have been less, as suggested by the
observations of Humphrey and Cope (1976). The
mean weight of adult males at Cadomin was signifi-
cantly greater (t-test, P<0.01) than that of juvenile
males, indicating that juveniles likely enter hiber-
nation weighing less than adults.
TABLE 4—Mean weights (+ 95% confidence intervals) of
Little Brown Bats captured at shelters in Edmonton and
Edson and at Cadomin Cave 19-20 September 1977.
Differences tested by Student’s t-test
Juvenile Juvenile Adult
Location females males males
Cadomin 8.7+0.4 (7) 8.6+0.4 (17) 9.8+0.2 (44)
Shelters 9:9) == O'S) (116) 92 == 056 0) O30 2518)
P<0.01 P<0.02 P<0.8
The earliest and latest dates that we have found
numerous bats hibernating were 20 September 1977
and 15 May 1975, respectively. Fifteen bats captured
on the spring date were all males, which suggests that
most or all females may have left the hibernaculum.
The sex ratio of hibernating bats favored males (Table
1) and was similar to that reported elsewhere
(Humphrey and Cope 1976).
The locations of the hibernacula of most Little
Brown Bats found in Alberta are unknown despite
inspection of mines and caves and continuing liaison
with caving organizations. The only suitable areas for
hibernacula within the province are in the Rocky
Mountains and a large area of karst in the northeast of
the province. The numerous coal mines of the central
and southern portion of the province, which would
likely offer potential hibernacula, have been routinely
blasted shut in accordance with provincial law. Three
hibernacula were located: Cadomin Cave and a cave
near Nordegg, both in the Rocky Mountains, over-
1979
winter between 500 and 1000 bats, and a third cave in
Wood Buffalo National Park in the northeast
contains a small number of hibernating Little Brown
Bats. Many bats would have to travel 450 km from
maternity roosts to reach either of these areas.
Reproduction
Of the 183 adult female Little Brown Bats collected
in maternity roosts prior to the onset of parturition
(15 June), 91% were pregnant. Twin fetuses were
observed twice among 312 pregnancies; 290 of the
single fetuses had implanted in the right horn. The
incidence of pregnancy among 162 bats that had
previously suckled young (95%) was significantly
higher (x test, P<0.001) than that among 21
unsuckled females (52%). Pregnant, unsuckled bats
generally weighed less than pregnant, suckled females
and they carried fetuses that often appeared smaller
than those of suckled bats. Three of eight unsuckled
bats caught in nurseries during mid-July were
resorbing embryos.
The earliest neonate was observed on 15 June
1975 and the latest date a pregnant bat was captured
was 15 July 1976. Parturition appeared to be highly
synchronous both within and between colonies as
reflected in the relatively consistent patterns of weight
gains of pregnant females and juvenile bats (Figures 2,
3) even though the samples were taken from colonies
in a variety of buildings and over a broad geographical
area. Based on fetal inspections and observations of
neonates, the majority of the bats born in 1972
through 1975 were born during the last week of June.
Parturition tended to commence earlier during 1976,
possibly owing to the early arrival of adult female bats
at the maternity roosts. Greater variation in fetal and
juvenile development was noted during that year.
Limited observations during 1977 indicated similarity
with 1976.
Like the females of many temperate-zone vesperti-
lionids (Asdell 1964), the majority of female M. /.
lucifugus in central Alberta do not bear young as
yearlings. Schowalter et al. (1978b) reported that
yearling bats comprised a smaller than expected
proportion of the 90 adult female bats in a nursery in
central Alberta and that all of the yearlings present
were non-parous. All eight of the unsuckled, non-
parous females from shelters that we have aged by
counts of dental annuli were young of the previous
year. Non-parous yearling bats probably comprise the
majority of the unsuckled bats found both in
maternity colonies and at shelters in central Alberta
(Table 2).
Juvenile Little Brown Bats are thought to be
reproductively active in New Mexico (O’Farrell and
Studier 1975) and Indiana (Humphrey and Cope
1976) as non-parous females were almost entirely
SCHOWALTER ET AL.: LITTLE BROWN BATS, ALBERTA
249
absent. Davis and Hitchcock (1965) found 11 of 21
yearling Little Brown Bats (aged by closure of finger
epiphyses) examined in New England to be pregnant,
suggesting that nonbreeding by juvenile females may
tend to increase with latitude. This may account for
the tendency for the increased proportion of females
in shelters in June with increased latitude (Table 3).
Differing timing of events both within the year and
within the life of individual female bats parallels
increased latitude. Later commencement of parturi-
tion in Little Brown Bats with increased latitude is
well documented (Fenton 1970; Humphrey and Cope
1976). O’Farrell and Studier (1975) have also
observed that the period of parturition becomes
abbreviated in more northern populations. Both these
trends are evident in comparisons of data from central
Alberta with those available from areas to the south.
In southern Alberta, which has a frost-free period
that is approximately 20 days longer than in central
Alberta (Government and University of Alberta
1969), parturition of M. 1. carissima is estimated to
extend from at least as early as mid-June to the first of
August, a much longer period than in M. /. lucifugus
in central Alberta. Determination of mean parturition
date of M. /. carissima was not possible from the
limited samples as parturition was highly variable
within and between colonies and between years;
however, parturition in 1975 and 1976 was later there
than in central Alberta. A collection made south of
Lethbridge on 12 July 1976 included seven lactating
and nine pregnant females, and two volant young.
Pregnant females were taken at the same colony 21
July 1975. Observations at this colony in July 1977
indicated that most young were born by at least mid-
July of that year. Visits to other colonies in the region
also indicated variability of parturition. The longer,
later, and more variable period of parturition in
southern Alberta may relate to minimal confrontation
with cold weather on the maternity range and to the
longer warm season which would, in many years,
allow late-born bats sufficient time to prepare for
hibernation.
O’Farrell and Studier (1975) related a brief parturi-
tion period of Myotis thysanodes to synchrony of
departure from hibernacula. Although it is possible
that most female Little Brown Bats found in central
Alberta follow this pattern, the geographic extent of
the region and the number of bats involved suggest
that this is not a complete explanation of the brief
parturition period observed. Female bats may return
to maternity roosts before food is abundant and
temperatures are favorable to take advantage of the
earliest arrival of favorable conditions. Low tempera-
tures are known to slow fetal development in some
bats (Racey 1973; Studier and O’Farrell 1972; Studier
et al. 1973) and fetuses may tend to be at a similar
250
stage of early development at the time of arrival of —
warm weather, regardless of the date of arousal from
hibernation and arrival at the colony of the females. In
years with mild early springs fetal development in
arriving females would not be slowed and an earlier,
more variable parturition period would result, as
observed in 1976. Whereas the gestation period of
early-arriving females may tend to be extended by
cool conditions in early May, gestation periods of
late-arriving females may be shortened by the com-
paratively high temperatures during long daylight
periods that are characteristic of late June. These
considerations suggest that the relative abbreviation
of the parturition period observed in Alberta, while
probably adaptive to bats living at higher latitudes,
may largely be a response to environmental condi-
tions rather than a specific regional adaptation of
bats.
The results described here were not obtained
through research designed specifically to examine
bat life-history. Detailed examination of a single
colony or local population rather than examination of
many colonies over a large geographical area would
clarify details of phenology and growth patterns.
Little Brown Bats occur far north of the present study
area and study of such boreal populations would
identify adaptations to exceedingly long daylight
periods. The trends identified here may be more
evident in those populations.
Acknowledgments
The research described here was part of a joint
program of the Alberta Fish and Wildlife Division,
ADRI(W) and the Veterinary Services Division of the
Alberta Department of Agriculture. Funding was
largely provided by the latter agency. The continued
interest and support of H. Vance and G. Whenham of
the Veterinary Services Division is gratefully ac-
knowledged. J. Bradley and W. Dorward of ADRI(W)
agreed to the use of data from their files and consented
to our use of rabies-suspect specimens. D. Bedard, H.
Boumans, P. Cole, W. Johnson, R. Masuda, D.
Meyer, D. Pipella, N. Previsich, B. Prins, and W.
Wynnyk assisted in various ways in field and
laboratory. H. Smith of the Provincial Museum of
Alberta provided bat bands and showed a continuing
interest in our research. Parks Canada and the
Alberta Parks Division consented to and assisted with
activities in parks. G. Lynch, A Todd, and J. Murie
reviewed drafts of this paper.
Literature Cited
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Banfield, A. W. F. 1974. The mammals of Canada. Univer-
THE CANADIAN FIELD-NATURALIST
Vol. 93
sity of Toronto Press, Toronto. 438 pp.
Davis, W.H. and H.B. Hitchcock. 1965. Biology and
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Received 10 November 1978.
Accepted 12 February 1979.
Winter Denning of the Striped Skunk in Alberta
JOHN R. GUNSON and RONALD R. BJORGE
Alberta Fish and Wildlife Division, O.S. Longman Building, 6909 116 Street, Edmonton, Alberta T6H 4P2
Gunson, J. R. and R. R. Bjorge. 1979. Winter denning of the Striped Skunk in Alberta. Canadian Field-Naturalist 93(3):
252-258.
Winter denning of the Striped Skunk (Mephitis mephitis) was studied in Alberta and adjacent Saskatchewan between 1971
and 1977. Most skunks denned under buildings and for periods up to 150 d. Skunks denned in communes and alone.
Communal dens contained a number of female skunks (mean of 5.8 per den) and usually one adult male skunk. Solitary dens
were occupied by males only, usually juveniles. Range of numbers of skunks in 61 communal dens was 2-19. Available
comparative data suggest commune size increases with latitude, an adaptation to optimize winter survival and reproductive
success. Age-specific selection seemed to operate in some communes with juvenile females excluded from certain dens.
Juvenile males visited den sites in fall, but only rarely denned with female communes.
Key Words: Mephitis mephitis, Striped Skunk, denning, communes, sex, age.
The Striped Skunk (Mephitis mephitis) dens for
extended periods during winter (Allen and Shapton
1942: Dean 1965; Verts 1967). More recent studies of
denned skunks during winter have explored the
potentials of disease transmission (Houseknecht
1969), activities (Sunquist 1974), physiological and
behavioral adaptations (Mutch and Aleksiuk 1977),
and habitat selection (Mutch 1977). We observed
winter denning of skunks during 1971-1977 in studies
of skunk rabies in Alberta and adjacent Saskatchewan
(Gunson et al. 1978) and in an intensive population
study of skunks in central Alberta (Bjorge 1977). This
paper summarizes observations of den descriptions,
numbers, sex and age, periods skunks spent in dens,
and of movements to and from dens, and considers the
adaptive significance of these aspects of winter
denning of skunks near the northern limit of the
species distribution.
Study Area
Most dens were located in the Alberta—Saskat-
chewan border area where the bulk of rabies-related
work was conducted (Figure 1). The area is aspen
parkland (Bird and Bird 1967) in the northern half and
prairie (Webb et al. 1967) in the southern half. It was
settled between 50 and 100 yr ago and current land use
has been in effect for many years. Cattle ranching and
dry-land farming predominate in the short-grass
prairie and cereal grain production and mixed-
farming in the parkland. The climate of the study area
is continental, characterized by warm summers and
cold winters (Government and University of Alberta
1969). Variations in annual precipitation and number
of frost-free days from north to south were = 28/18 cm
and = 100/130 d.
The 130-km? intensive study area (Tofield), located
= at 53°23’N, 112°43’W, appeared typical of the aspen
parkland. Eight major forested areas (mostly Trem-
bling Aspen, Populus tremuloides), ranging 30-65 ha,
occurred as well as numerous smaller areas with trees
and brush along fence rows, road allowances, farm-
yards, and wetlands. About 150 ponds, many of them
seasonal, dotted the area. Predominant crops were
cereal grains, forage, livestock, and poultry. Of 112
farmyards on the study area, 40 were abandoned.
Methods
Dens were located during six winters (1971-72 to
1976-77) primarily by visual examination of likely
locations such as old buildings, culverts, and ground
burrows. When located, skunks were removed from
dens chiefly by carbon-monoxide gassing with two 2-
cycle gasoline-operated motors, shooting (22-calibre
rifle), and kill-trapping with mostly 220 Conibear
traps. Collection of all skunks was often possible
through removal of floorboards or occasionally by
jacking or digging under foundations. Only dens
- examined during November to March, and only
those dens where collectors were convinced all skunks
were captured, were used in calculations of numbers
and of sex and age.
Skunks were live-captured at Tofield during the
non-winter period by trapping (National, Rudolph
Skunker, and wooden and metal box traps were used),
night-lighting (Jacobson 1969), and excavation of
maternal dens. Live-captured skunks were handled
according to the techniques of Jacobson et al. (1970)
including netting, ear-tagging, weighing, sexing, and
release. Radiotransmitters, similar to those used by
Brand et al. (1975) and developed by personnel of the
Department of Electrical Engineering, University of
Alberta, Edmonton, were fitted to skunks. Signals
were received via a truck-mounted non-directional
whip antenna or a hand-held directional loop antenna
and a portable receiver. Locations were determined by
triangulation.
DS)
1979 GUNSON AND
BJORGE: STRIPED SKUNKS WINTER DENNING
U3)3)
| ALBERTA
N
100 km
|
62 mi x
Edmonton
ofield
Parkland
= ~
f
7
Calgary
\
Wen f
\ e
\ e
eye Pra
\ 7 ee “e
Ficure |. Locations of 61 communal dens of Striped Skunks in prairie and parkland portions of Alberta and adjacent
Saskatchewan.
Dental ages of collected skunks were determined by
counts of cemental annuli in canines (Rakowski 1972;
Casey and Webster 1975) after histological prepar-
ation at Matson’s Audiovisual and Microscopic,
Milltown, Montana. Live-captured young of the year
were classified as juveniles until 31 December on the
basis of weight, nipple development, and general
appearance.
Results and Discussion
Location and Description of Dens
Our searches for winter dens were directed prima-
rily at rural, usually abandoned, buildings; thus the
precise proportional use of buildings versus burrows,
culverts, and other possible locations during winter
was not determined. Of 73 communal ( >1 skunk)
dens, 72 were located under buildings; one was in a
culvert. Of 44 solitary (1 skunk) dens, 37 were under
buildings, five were in culverts, and two were in
ground burrows. Five of the six culvert dens were
located in the southern prairie region. Both ground
burrows were located at Tofield after extensive
searching and tracking in snow during early winter
1971-72. The relatively large number of female
skunks (see next section) under some buildings and
the availability of other buildings without skunks, but
used during previous winters, suggested that most
female skunks, especially in the parkland, denned
under buildings.
During a depopulation at Tofield in December
1974, 19 (53%) of 36 female skunks tagged during
June to October of that year were removed from six
communal dens. Depopulation was incomplete at one
den and skunks could not be removed from one other
potential communal den because of its concrete
structure. The above, and probable losses from death
and emigration between summer and winter, support
our belief that the majority of female skunks in the
parkland den under buildings. Comparable recapture
rate at buildings for males was only 11.9%, suggesting
that many male skunks den in locations other than
buildings. Of the two ground burrow dens located,
one was used by a tagged male; the sex of the skunk in
the other burrow was not determined.
Related research (Gunson et al. 1978) implied
relatively heavy skunk use of buildings during winter
in the parkland versus prairie. Skunk depopulations
by poisoning at buildings effectively reduced skunk
populations in the parkland, but not in the prairie.
Use of buildings as winter dens of skunks appeared
greater in our study area than that recorded in more
southerly areas. Allen (1939) and Storm (1972)
reported that buildings were used only occasionally by
skunks in Michigan and Illinois, respectively, It may
254
be that buildings were more available to skunks in our
area than in other areas. Larger commune size and
lower densities (unpublished data) of skunks in our
area compared to more southerly areas probably
favored the selection of building dens. A preference
for building dens would result in greater proportions
of skunks utilizing such dens in areas or periods of
lower density. Building dens offer skunks more space
for walking and use of latrines and for breeding during
years of longer winters.
Skunks made use of the space between ground and
floorboards in most building dens. All winter dens
that we observed contained dried vegetation, mostly
grass. Ground burrows, usually associated with
cellars, were present under some buildings. Construc-
tion of burrows was likely necessary in some cellars to
create a more confined den area.
Numbers, Sex, and Age of Skunks in Dens
A mean of 6.7 skunks was collected from 61
communal dens where collectors were convinced all
skunks were taken (Table 1). Females were more
common incommunal dens than males; in fact, no den
contained more than one live male. Allen (1939) noted
that only one male was present with denned females in
his area. We obseved 43 dens of solitary males;
denning by solitary females was not observed. Dens of
solitary males and females were reported by Allen
(1939) and Allen and Shapton (1942) in Michigan.
TABLE 1—Composition of 61 communal winter dens of the
Striped Skunk in Alberta and adjacent Saskatchewan (dens
where all skunks were collected)
Number of skunks
Sex
Total Mean (range)/den
Females 353 5.8 (1-18)
Males 53 0.9 (O- 1)
Both sexes 406 6.7 (2-19)
Somewhat more skunks were taken from communal
dens in parkland versus prairie (means of 6.8 of 32
dens and 6.4 of 29 dens, respectively (Student’s t-test,
P<0.4)). The largest commune was located in
Township 57, the most northerly den we observed.
Comparable data from other areas, although meagre,
suggest commune size increases with latitude. Shirer
and Fitch (1970) working in Kansas noted three
skunks denned together, but for only short periods
during winter. Numbers of skunks in dens was 2 to 11
in 10 dens (mean of 5.1) in Michigan (Allen 1939;
Allen and Shapton 1942), two to six per den in
Minnesota (Houseknecht 1969; Sunquist 1974), and
THE CANADIAN FIELD-NATURALIST
Vol-93
eight in one den in Colorado (Yeager and Woloch
1962).
Larger communes in more northerly areas are likely
an adaptation of the species to more rigorous
climates. Mutch and Aleksiuk (1977) demonstrated
that den temperatures vary directly with numbers of
skunks in the den. It follows that females in larger
groups would expend less energy in surviving longer
winters, during which they do not feed, and conserve
more energy for reproduction. Length of winter
importantly influences production of skunks. Litters
of young skunks at Tofield were significantly larger
(Student’s t-test P<0.05) in 1973 (mean of 5.2) after a
relatively mild winter compared to size of litters
following the severe winter of 1973-74 (mean of 2.5)
(Bjorge 1977).
Juvenile (<1 year) females represented 47.7% of
327 aged females (Figure 2). In contrast, juvenile
70
60
50 Solitary Dens
Males
40 n=38
0
20
10
50 Communal Dens
Males
n=51
50
is)
PROPORTION (°%o) OF SAMPLE
Communal Dens
Females
n=327
0
O
; : ;
| =] = —
0.5 15 2.5 3.5 4.5 5.5 6.5
AGE-GROUP
FIGURE 2. Age-structure of Striped Skunks in solitary and
communal dens in Alberta and adjacent Saskatchewan.
1979
males comprised only seven (13.7%) of 51 aged males
in communal dens. Most of the lone males were
young; 26 (68.4%) of 38 aged solitary males were
juveniles. No lone male was over 3 yr of age.
Comparisons of age structures of skunks in pop-
ulations during summer and fall and in communal
winter dens at Tofield (Table 2) indicated that the
juvenile proportion in the female cohort was some-
what similar in dens and free-ranging skunks. Juvenile
males, however, although common (> 60%) in male
cohorts in populations, were absent in six communal
dens during winter. In summary, winter dens of the
Striped Skunk in our area included communal
denning of females, usually with one adult male, and
solitary denning of other males, including most
juveniles.
TABLE 2—Comparisons of age structures of Striped
Skunks in six communal dens and in populations on the
Tofield, Alberta study area
Proportion (%) juveniles (n)
Source Period
Male Female
Communal dens 1974-75 O0( 5) 47 ( 32)
Population! 1974 66 ( 41) 35 ( 34)
Population! 1971-74 62 (134) 37 (140)
'Live-trapped during June through October; excludes
captures at communal dens in September and October.
Den Formation
Live-trapping at Tofield during September through
October 1971-74, indicated that male skunks visited
winter den sites other than the one they would occupy.
From two to three male skunks were captured at each
of six sites at which communes were forming during
fall. Because only one male winters at each commune
a male selection process must occur. Much of the
selection might be a result of avoidance, probably
juveniles tending to avoid more experienced adults.
At each of two sites in 1974, adult and juvenile males
were captured at the den site in fall; only the adults
were present with the female commune in winter.
Some competition, involving fighting, probably oc-
curs during the fall pre-denning period. Three males
with fresh scars on ears and face were observed during
that period.
Some unsuccessful (solitary) males visited com-
munal dens during winter. A farmer poisoned a male
skunk at one communal den in southern Alberta in
late February. During the first week of March four
live skunks, including one male, were collected at the
den. We believe it highly probable that the second
GUNSON AND BJORGE: STRIPED SKUNKS WINTER DENNING
DES
male joined the commune after the death of the former
male. Tracks of skunks to and from dens were
common during that collection period. At each of two
other communal dens one live and one dead male
skunk were collected. During a thaw in February
1972, at Tofield, a solitary male visited a commune
1 km from his burrow.
Data from Tofield suggested that occupation of a
specific communal den by females in at least some
cases probably depended on age and experience. Five
of seven females at one winter den at Tofield were
adults, only one of which was a yearling. R.R.B.
considered this area to have a higher density of skunks
than most of the remainder of the Tofield area. At
least four of the eight skunks at that winter den were
captured and tagged during 1973 and 1974 at another
den site used the previous three winters, but not in
1974-75. All or most of the group moved some 2.5 km
to the new winter den during the fall pre-denning
period in 1974. An adult male wintered alone at the
original den. These observations suggest the com-
mune was highly cohesive, and may explain the
observed relatively low proportion of juveniles. Other
young females may have been forced to den elsewhere.
We noted that two other communes changed dens
between years.
Another communal den of seven females, located in
an area R.R.B. considered to be of lowskunk density,
consisted of six juveniles and one yearling. We
collected some evidence that suggested these were not
siblings. First, as discussed later, juvenile females at
Tofield were highly mobile, travelling many kilo-
metres to communal dens (Bjorge 1977). If such
extensive dispersal of juvenile females is common it is
unlikely that siblings would den at the same location.
Second, of 62 young skunks, including 27 females,
tagged in natal dens in 1973 and 1974, none was
recaptured in communal dens on the study area even
though survival of young was demonstrated at seven .
of 15 dens. Survival was not recorded at the other
natal dens. Allen (1939) concluded that winter groups
of female skunks in his area were not families.
Juvenile proportion of females in 56 communes was
not bimodally distributed (Figure 3) as one might
have expected from the selection process outlined
above. Selection against juveniles at some dens could
potentially create a situation of most dens with either
low or high proportions of juveniles. Thirty-three
(59%) of 56 dens contained < 30% or > 60% juveniles.
Many of the dens in our sample were located in areas
where skunks were being depopulated for rabies
control. Because skunk sign was more abundant and
more readily observed at communal dens than at
solitary dens, skunks were more likely to be discov-
ered and poisoned at communal dens. Higher than
normal mortality at communal dens may account for
256
W
Number of Dens
u
0 11621 0«631)S 41—s“O51 4 vi at n
| I | | I
10 20 30 40 50 60 70 80 90 100
Juvenile Proportion
%
FIGURE 3. Juvenile proportion of female Striped Skunks in
56 communal dens in Alberta and adjacent
Saskatchewan.
the variation between observed and expected distri-
butions. Juveniles might be more readily accepted
into dens of reduced size.
Denning Period
Success of live-capture at sites of communal dens
at Tofield increased from 69 and 124/ 1000 trap-nights
in July and August, respectively, to 275 in September;
skunks began to harvest vegetation around den
entrances in late September, as observed by Sunquist
(1974) in Minnesota. This seemed to represent a pre-
denning activity as skunks were still free-ranging
during September and October, as evidenced by
success of night-lighted captures along roads (Gunson
et al. 1978) and by live-capture success at non-denning
sites at Tofield. Adult skunks likely began denning
before juveniles. Night-lighted captures of free-
ranging skunks during late September and October
were highly biased towards juveniles (unpublished
data), and mean distances of adult females from
communal dens at Tofield during September and
October were significantly (Student’s t-test P< 0.05)
less than for juveniles. Live-capture at communal den
sites was most successful in October (306/ 1000 trap-
nights). Sunquist (1974) noted that juveniles were
more active than adults in November and initiated
THE CANADIAN FIELD-NATURALIST
Vol. 93
winter denning an average of 7 d later than adults. In
1973 activities of skunks beyond the immediate
vicinity of winter dens at Tofield were terminated bya
25-cm snowfall on 10 November.
Observations at 10 occupied dens during winter
1971-72 indicated that activity beyond a few metres of
the den did not occur during early December to mid-
February. A marked increase of activity was evident
at six sites following a thaw on 12 February, with
tracks leading from or to dens in four instances.
Activity during mid-winter thaws was common
during the study period, but evidently was mostly
confined to males; of 13 road-killed skunks examined
during December through February, 12 were males.
Dean (1965) also noted the reactivation of winter-
denned skunks during above-freezing temperatures.
Emergence of skunks from two ground burrows
occurred in mid-February and mid-March 1972. The
latter emergence occurred in an area of snow
accumulation and the emerging skunk had to burrow
through approximately 50 cm of snow to leave the
den. Movements of skunks from dens were common
by mid-March. Males appear to leave dens earlier
than females. Two skunks captured away from dens at
Tofield in March were males. Preponderance of male
skunks in fur harvest records (see Verts 1967) are
likely explainable by greater and earlier activities of
male skunks during winter and spring as recorded
here.
Dates of departure of six radio-collared adult
female skunks from dens at Tofield were all between 2
and 9 April 1974. Because some of the six skunks may
have denned before the 10 November snowfall (day 1),
minimum period in winter dens during 1973-74, a
winter of unusual severity, was 143 to 150 d. During
the winter of 1971-72, a winter of moderate severity,
most skunks were dénned by mid-November and
skunks in communes were mostly inactive until mid-
March, a period of approximately 120 d.
These observations indicate that female skunks
generally occupy winter dens in the parkland of
Alberta between 120 and 150 d, and that males usually
den for shorter periods. In Alberta the winter period is
shorter in the prairie portion of the province and
skunks in that area denned for correspondingly
shorter periods. We observed numerous tracks
leading from dens in southern areas of the province
during March and male-female pairs of skunks were
often observed at non-winter sites during that month.
Skunks occupied winter dens 79-125 d in Minnesota
(Sunquist 1974) and 62-87 d in Illinois (Verts 1967;
Storm 1972).
Movements to and from Winter Dens
Juveniles travelled further than adults to winter
dens; mean greatest distances from dens of 47 adult
1979
and 22 juvenile skunks during summer were 2.3 and
3.1 km, respectively. Adults tended to occupy the
same general areas during consecutive summers
(Bjorge 1977). Four adult females, monitored by
telemetry from 46 to 85 d after leaving the den in
spring, travelled greatest straight-line distances of 1.6
to 3.4 km. Two other females, monitored from July
and August travelled 3.0 and 3.5 km to their winter
dens.
Movements of juvenile female skunks to winter
dens were 13.7, 5.9, and 5.6 km from initial locations
in July, September, and October, respectively. One
other juvenile female travelled 21.7 km to a farmyard
where she was shot before denning. Extensive
dispersals of juveniles were recorded in another
intensive study area in the prairie region in Alberta (P.
Andersen, University of Calgary, personal commun-
ication). The observed mobility of juveniles may be
related to a search for acceptance into a winter
commune.
Concluding Discussion
Carnivores have evolved a wide range of species-
specific winter denning strategies varying from the
mostly solitary denning of bears to the highly
communal denning of female Striped Skunks as
reported here. Social organization of denning would
appear to be importantly related to reproduction in
carnivores. European Badgers (Meles meles) and
American Badgers (Taxidea taxus) apparently den
either in pairs or solitarily (Neal 1948; Ewer 1973;
Lindzey 1978). Breeding of badgers and bears occurs
during summer (see Ewer 1973), and is followed by
delayed implantation; thus, communal denning in
these species would not enhance reproduction.
Because Striped Skunks breed during or immediately
after winter denning, reproduction is potentially more
successful where numbers of females are available to
at least one male. Ewer (1973) noted that Verts
observed one male would breed several captive female
skunks in short order; such behavior would ensure
that all females become pregnant. Pregnancy rates of
Striped Skunks in Alberta were high during the study
period (unpublished data).
Communal winter denning is common in Raccoons
(Procyon lotor), although group size is usually small
(Ewer 1973), despite one den of 23 individuals (Mech
and Turkowski 1966). Family groups of Raccoons
den together (Whitney and Underwood 1952). Our
observations suggest that family groups of Striped
Skunks in Alberta do not often den together. Genetic
variation would be maintained by family separation in
areas where a significant proportion of breeding
occurs during winter denning.
In summary, communal denning appears to have
adaptive value for winter survival and reproductive
GUNSON AND BJORGE: STRIPED SKUNKS WINTER DENNING
a3}
success of Striped Skunks, especially in more
northern latitudes where the period of inactivity and
fasting is longer, where energy conservation can be
critical, and where breeding may be influenced by
winter distribution.
Acknowledgments
This study was supported by the Veterinary
Services Division, Alberta Department of Agriculture
and by the Fish and Wildlife Division, Alberta
Department of Recreation, Parks and Wildlife. The
Saskatchewan Department of Natural Resources
(now Tourism and Renewable Resources) kindly
provided permission to collect skunks in adjacent
Saskatchewan. The capable technical assistance of P.
Andersen, P. Cole, W. Etherington, E. Ewaschuk, W.
Johnson, L. Pecharsky, D. Pipella, D. Schowalter,
and W. Wynnyk is acknowledged. D. Schowalter
kindly reviewed earlier drafts of this paper. W. M.
Samuel of the University of Alberta supervised much
of the Tofield studies during 1973 and 1974.
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Casey, G.A. and W.A. Webster. 1975. Age and sex.
determination of Striped Skunks (Mephitis mephitis)
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Dean, F. C. 1965. Winter and spring habits and density of
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census to Striped Skunk population studies. M.Sc. thesis,
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Pp:
258
Jacobson, J. O., E. C. Meslow, and M. F. Andrews. 1970.
An improved technique for handling Striped Skunks in
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510-512.
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northwestern Utah. Journal of Wildlife Management
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Striped Skunks in Delta Marsh, Manitoba. Canadian
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radio-tracking of movements and denning habits of the
THE CANADIAN FIELD-NATURALIST
Vol. 93
Raccoon, Striped Skunk and Opossum in northeastern
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Received December 1978
Accepted 5 March 1979
Range Extensions of Vascular Plants in Northern Yukon Territory and
Northwestern District of Mackenzie
JOHN A. NAGY,! 3 ARTHUR M. PEARSON,! 4 BERNARD C. GOSKI,! 5 and WILLIAM J. Coby?
1Canadian Wildlife Service, Environment Canada, Edmonton, Alberta T5K 2J5
2Biosystematics Research Institute, Canada Agriculture, Ottawa, Ontario KIA 0C6
3Present Address: Lethbridge Community College, Lethbridge, Alberta TIK 1L6
_ 4Present Address: Government of the Yukon Territory, Whitehorse, Yukon Y1IA 2C6
5Present Address: Alberta Fish and Wildlife Division, Edson, Alberta TOE OPO
Nagy, John A., Arthur M. Pearson, Bernard C. Goski, and William J. Cody. 1979. Range extensions of vascular plants in
northern Yukon Territory and northwestern District of Mackenzie. Canadian Field-Naturalist 93(3): 259-265.
Between 1970 and 1975 plant communities in the Yukon Territory and the District of Mackenzie north of 67°N were
investigated. We record three taxa new to the known flora of the Yukon Territory and one new to the flora of the District of
Mackenzie. Among the other vascular plants listed here, nine are new to the range predicted by Hultén within the Yukon
Territory, 23 are extensions within predicted ranges and thus corroborate these predictions, and 29 are extensions beyond
predicted ranges.
Key Words: vascular plants, Yukon Territory, District of Mackenzie, flora.
Between 1970 and 1975 over 800 collections of — tics Research Institute, Canada Agriculture, Ottawa,
vascular plants, representing 220 species, 53 sub- Ontario.
species and seven varieties, were made during Cana-
dian Wildlife Service studies in the northern Yukon Collection Sites
Territory. Hultén (1968), Welsh and Rigby (1971), Approximate collection site locations are shown in
and Wein et al. (1974) have reported extensive Figure 1. Other site data are listed below.
collections and recent range extensions for vascular 1) Sam Lake, 68°25’N, 138°35’W, elevation ca.
plants from our study area. We record here additional 427 m (1400 ft). Marshy polygonal; dry-to-wet
species to the reported flora of the Yukon Territory tundra and hummocky lakeside areas; marshy
and northwestern District of Mackenzie. We also note and wet shrubby lakeside areas.
species that occur outside of the ranges predicted by 2) Canoe River, 68°37’N, 138°43’W, elevation ca.
Hultén (1968), or were collected for the first time in 305 m (1000 ft). Wet sedge meadows west of
northern Yukon Territory within ranges predicted. In river.
addition, a number of species are recorded as the 3) Ridge 1.6km east of Sam Lake, 68°24’N,
northernmost or southernmost collections within 138°33’W, elevation ca. 488 m (1600 ft). Dry
ranges from which collections are known. rocky and moist-to-wet tundra on ridge; dry
The study area and collection locations are shown south- and north-facing scree slopes; wet snow
in Figure 1. The 31 collection sites from which plant bed community near drainage.
specimens were obtained ranged in elevation from 122 4) Anker Creek, 68°43’N, 137°33’W, elevation ca.
to 152 masl on the Arctic Coastal Plain, 244 to 122 m (400 ft). Alder-willow bluffs on moist-to-
427 masl on the Old Crow Flats, and 152 to wet south-facing slope; dry rocky talus slope.
1067 masl in the Barn, British, and Richardson 5) Trail River, 68°57’N, 138°56’W, elevation ca.
mountains. Bostock (1961) and Welsh and Rigby 152 m (500 ft). South-facing river bank.
(1971) describe the physiography of the area. 6) Headwaters of Anker Creek, 68°30’N, 138°19’W,
Our activities were primarily concentrated in areas elevation ca. 945 m (3100 ft). Alpine slope and
that had not received attention from previous investi- tundra.
gators. Collectors were A. M. Pearson 1970, 1973, 7) 6.4km northwest of Sam Lake, 68°28’N,
1974, 1975; J. A. Nagy 1974, 1975; M. Dennington 138°40’W, elevation ca. 457 to 762 m (1500 to
1975; B. C. Goski 1973, 1974: D. Pearson 1974; C. B. 2500 ft). Areas of dry, rocky and moist-to-wet
Larsen 1973, 1974; J. W. Nolan 1973. The collections tundra; moist-to-wet and swampy areas in and
made during 1970 and 1973 were identified by A. E. along edge of White Spruce ( Picea glauca) stand;
Porsild, National Museum of Natural Sciences, willows (Salix spp.) in and along edge of White
Ottawa, and those collected during 1974 and 1975, by Spruce stand; grassy slopes on upper edge of
W. J. Cody and G. A. Mulligan (Draba), Biosystema- White Spruce stand; dry rocky ridge and slope.
259)
260 THE CANADIAN FIELD-NATURALIST Vol. 93
0 kilometres ©O
BEAUFORT SEA
SS
Oo miles 30
Herschel
MACKENZIE
BAY
68°
{ «e eg
40° (38° 136°
FicurE |. Locations of 31 vascular plant collection sites in the northern Yukon Territory and northwestern District of
Mackenzie.
1979
8)
9)
10)
11)
12)
13)
14)
15)
16)
17)
18)
19)
20)
21)
22)
23)
24)
25)
26)
2i))
Ridge 1.6km (1 mi) north of Sam Lake,
G8S225UNe 1382350W, elevation’ “cae. 58m
(1700 ft). Dry rocky slopes.
Dog Creek, 68°25’N, 138°38’W, elevation ca.
427 m (1400 ft). Willows along drainage; and
tundra.
Headwaters of Boulder Creek, 68°26’N,
138°13’W, elevation ca. 488 m (1600 ft). Moist-
to-dry rocky south-facing slopes; stabilized
south-facing talus slope; mossy wet stream bed;
moist tundra along creek.
Ridge above Boulder Creek, 68°25’N, 138°16’W,
elevation ca. 914 m (3000 ft). Dry rock south-
facing slope.
Tributary of Babbage River, 68°38’N, 138°01’W,
elevation ca. 427 m (1400 ft). Wet mossy stream
bank; wet streamside willow stand; dry south-
facing slope.
Tributary of Fitton Creek, 68°33’N, 138°10’W,
elevation ca. 457 m(1500 ft). Moist drainage; dry
rocky ridge.
Trail River, 68°58’N, 138°58’W, elevation ca.
396 m(1300 ft). Moist drainage; dry south-facing
slope.
Spruce Creek, 68°56’N, 138°41’W, elevation ca.
305 m (1000 ft). White Spruce stand on dry
south-facing slope.
Old Crow Flats, 68°09’N, 139°35’W, elevation
ca. 274 m (900 ft). Bog.
Old Crow Flats, 68°12’N, 139°49’W, elevation
ca. 305 m (1000 ft). White Spruce stand on dry
ridge; marshy area.
Old Crow Flats, 68°12’N, 139°48’W, elevation
ca. 305 m (1000 ft). Sphagnum bog.
Old Crow Flats, 68°01’N, 139°48’W, elevation
ca. 244 m (800 ft). Sphagnum bog.
Headwaters of Babbage River, 68°38’N,
139°32’W, elevation ca. 457 m(1500 ft). Willows
along stream bank; tundra.
Margaret Lake, 68°50’N, 140°36’W, elevation
ca. 457 m (1500 ft). Tundra along Firth River.
Babbage Falls, 68°43’N, 139°03’W, elevation ca.
274 m (900 ft). Rock crevices along falls; dry
rocky south-facing slope.
Bell River, 67°43’N, 136°33’W, elevation ca. 427
to 457 m (1400 to 1500 ft). Scree slope; White
Spruce — White Birch (Betula papyrifera) —
willow stand; sphagnum bog; moist slope.
Firth River, 69°20’N, 139°33’W, elevation ca.
152 m (500 ft). North-facing rock outcrop.
Boulder Creek, 68°25’N, 138° 15’W, elevation ca.
457 m (1500 ft). Moist creek-side areas near
headwaters.
Sam Lake, 68°30’N, 138°35’W, elevation ca.
762 m (2500 ft). Dry hilltop.
Mount Sedgwick, 68°58’N, 139°01’W, elevation
NAGY ET AL.. NORTHERN VASCULAR PLANTS
261
ca. 305 m (1000 ft). Dry slope near ridge top;
moist creek-side area.
Old Crow, 67°35’N, 139°50’W, elevation ca.
250 m (820 ft). Roadside near airstrip.
Rapid River, 68°16’N, 137°08’W, elevation ca.
914m (3000 ft). Sedge association in thick
organic soil on moist south-facing rock slide.
Margaret Lake, 68°48’N, 140°36’W, elevation
ca. 457 m (1500 ft). Lakeside sphagnum bog.
Bear Creek, 67°58’N, 136°12’W, elevation ca.
1067 m (3500 ft). Moist basin.
28)
29)
30)
31)
Vascular Plant Range Extensions
The 65 taxa reported here are listed with collection
sites, habitats, collection years, and collection
numbers. Nomenclature follows Hultén (1968) and if
names used by Porsild and Cody (1968) differ, they
are also given, in parentheses. A complete set of
voucher specimens has been deposited in the Cana-
dian Wildlife Service Herbarium at Edmonton, with
duplicates of the 1974 collections deposited at the
Canada Agriculture Herbarium in Ottawa (DAO).
Three taxa, Luzula Wahlenbergii ssp. Wahlen-
bergii, Potentilla rubricaulis, and Galium Brandegei
are reported as new to the flora of the Yukon
Territory, and one species, Cerastium maximum, as
new to the flora of the Continental Northwest
Territories.
The new range data provided here on the taxa listed
represents a contribution to our knowledge of the
phytogeography of these taxa, as well as a contribu-
tion towards a flora of Yukon Territory.
Lycopodiaceae
Lycopodium annotinum ssp. pungens (Stiff Club-moss).
Site 1: shrubby lakeshore, 74-58; site 4; alder, 74-189; site 5:
river bank, 74-10; site 23: sphagnum bog, 74-683. Helps
complete gaps in the predicted distribution between sites in
northwestern District of Mackenzie and Alaska, extending
the known range northward to the Arctic Coastal Plain
(Hultén 1968).
Lycopodium complanatum (Ground-Cedar). Site 23: mixed
White Spruce — birch — willow, 74-68]. Extends the known
range northward from collection localities reported by
Hultén (1968) and Wein et al. (1974) along the Yukon River
drainage.
Polypodiaceae
Cystopteris fragilis ssp. fragilis (Fragile Fern). Site 22:
crevices along falls, 74-675. Hultén (1968) predicted the
occurrence of both ssp. fragilis and ssp. Dickieana in
northern Yukon Territory and did plot one collection from
Yukon Territory north of 65°N, along the Yukon River
drainage near the Alaska border. Welsh and Rigby (1971)
reported three specimens from the British Mountains, but
did not give their collections a subspecific designation. The
specimen reported here is the first record of ssp. fragilis
262
within the predicted range for northern Yukon Territory
(Hultén 1968).
Dryopteris fragrans (Fragrant Cliff-Fern). Site 5: river bank,
74-13; site 14: south-facing slope, 74-435. Welsh and Rigby
(1971) reported the first collection from the northern Yukon
Territory. The specimens reported here are the most
northern collections, and help complete gaps in the predicted
distribution for northern Yukon Territory (Hultén 1968).
Sparganiaceae
Sparganium minimum (Bur-reed). Site 18: emergent
aquatic, 74-556. Extends the known range significantly
northward from collection localities in west central Yukon
Territory to the Old Crow Flats (Hultén 1968).
Potamogetonaceae
Potamogeton alpinus ssp. tenuifolius. Site 18: sphagnum
bog, 74-550. Extends the known range to the northwest of
Welsh and Rigby’s (1971) recent first collection for Yukon
Territory north of 65°N.
Potamogeton perfoliatus ssp. Richardsonii (P. Richard-
sonii) (Red-head Pondweed). Site 16: submerged aquatic,
74-511. First record from the predicted range for the Yukon
Territory north of 65° N (Hultén 1968). Helps bridge the gap
in the distribution between collection sites in Alaska and
northwestern District of Mackenzie.
Gramineae
Calamagrostis canadensis ssp. Lansdorffii (Blue-joint).
Site 18: sphagnum bog, 74-549; site 25: creek bank, 73-/,
73-3. Hultén (1968) predicted the occurrence of both ssp.
canadensis and ssp. Langsdorffii in northern Yukon and
reported collections of both subspecies adjacent to the
Yukon Territory border. Welsh and Rigby (1971) reported a
specimen of C. canadensis from the Old Crow Flats, but did
not give their collection a subspecific designation. This is the
first report of ssp. Langsdorffii from the predicted range for
Yukon Territory north of 65°N.
Calamagrostis purpurascens. Site 21: tundra, 74-667. Hultén
(1968) cited collections from adjacent areas in Alaska and
northwestern District of Mackenzie, while Welsh and Rigby
(1971) reported the first collections from northern Yukon
Territory. The specimen reported here is the most northern
collection within the predicted range for Yukon Territory
(Hultén 1968).
Festuca ovina ssp. alaskana. Site 8: dry rocky ridge, 74-320.
Hultén (1968) reported a collection from the Yukon River
drainage near the District of Mackenzie border, and Cody
has collected it in the Richardson Mountains to the east in
District of Mackenzie (Cody and Porsild 1968). The
specimen reported here is the most northern collection in the
predicted range in Yukon Territory.
Cyperaceae
Carex glacialis. Site 7: rocky south-facing slope, 74-254.
First record within the predicted range for Yukon Territory
north of 65°N (Hultén 1968).
Carex lapponica (C. canescens var. subloliacea). Site 19:
sphagnum bog, 74-560. Extends the known range signifi-
cantly northward from central Yukon Territory to the Old
Crow Flats (Hultén 1968).
THE CANADIAN FIELD-NATURALIST
Vol. 93
Carex membranacea. Site 6: alpine tundra, 74-346; Site 7:
White Spruce stand, 74-615; site 7: dry rocky ridge, 74-288;
site 20: tundra, 74-586a. Hultén (1968) plotted collection
localities from adjacent areas in Alaska and District of
Mackenzie. The specimens reported here are the first
collections from the predicted range for Yukon Territory
north of 65°N.
Carex Williamsii. Site 16: bog, 74-508. First reported for
northern Yukon but is from south of the range predicted by
Hultén (1968) for the northern Yukon Territory, and
represents only the fourth collection for Yukon Territory
(Porsild 1975).
Eriophorum angustifolium ssp. triste (E. triste). Site 6: alpine
tundra, 74-347; site 7: dry rocky ridge, 74-118. Hultén (1968)
indicated a collection from the Firth River drainage in
Alaska near the Alaska border. The specimens reported here
are the first collections from the predicted range for Yukon
Territory north of 65°.
Eriophorum russeolum var. albidum. Site 1: marsh, 74-75.
Hultén (1968) cited collection sites from adjacent areas in
Alaska and northwestern District of Mackenzie. The
specimen reported here is the first collection from the range
predicted by Hultén (1968) for northern Yukon Territory-
Porsild (1951, 1975) has reported it from along the Canol
Road and the Ogilvie Mountains.
Araceae
Calla palustris (Wild Calla). Site 18: sphagnum bog, 74-546;
site 28: roadside, 73-8. Hultén (1968) reported a collection
from the headwaters of the Yukon River. Our collection
extends the known range northwestward to the Old Crow
Flats.
Lemnaceae
Lemna trisulca (Star-Duckweed). Site 18: floating aquatic in
sphagnum bog, 74-554. Northernmost collection within the
range predicted by Hultén (1968) for Yukon Territory.
Juncaceae
Juncus articus ssp. alaskanus (J. balticus var. alaskanus).
Site 21: riverside tundra, 74-664. Welsh and Rigby (1971)
reported the first collection (presumably var. alaskanus) of J.
arcticus from northern Yukon Territory, but did not give
their specimen a subspecific designation. This is thus only the
second record of ssp. alaskanus within the range predicted by
Hultén (1968) for Yukon Territory north of 65°N.
Luzula arctica (L. nivalis). Site 10: south-facing slope, 74-
170. First report from the predicted range for Yukon
Territory north of 65°N (Hultén 1968) and helps bridge the
gap between collection localities in Alaska and northwestern
District of Mackenzie.
Luzula arcuata s. lat. Site 13: drainage, 74-383. First
collection from Yukon Territory north of 65° N, and is from
near the northern limit as predicted by Hultén (1968).
Luzula multiflora s. lat. Site 3: dry rocky hillside, 74-633; site
7: tundra, 74-24]. First collections within the range predicted
by Hultén (1968) for Yukon Territory north of 65°N.
Luzula parviflora. Site 12: stream bank, 74-2/2. First
collection from Yukon Territory north of 66°N, extending
the known range northward to the Barn Mountains (Hulten
1968).
1979
Luzula Wahlenbergii ssp. Wahlenbergii. Site 1: polygonal
ground, 74-269. Hultén (1968) reported a collection from
northwestern District of Mackenzie adjacent to the Yukon
Territory border. The specimen reported here is the first
collection from the range predicted by Hultén (1968) for
Yukon Territory and is new to the Territory.
Orchidaceae
Spiranthes Romanzoffiana (Hooded Ladies’-Tresses). Site
7: bog, 74-503; sphagnum bog, 74-56]. First collection
reported from Yukon Territory north of 65° N, extending the
predicted range westward from collection localities in
northwestern District of Mackenzie to northern Yukon
Territory (Hultén 1968).
Salicaceae
Salix Barrattiana. Site 14: drainage, 74-463. First specimen
collected in northern Yukon Territory, extending the
predicted range eastward from collection sites in Alaska
(Hultén 1968).
Caryophyllaceae
Cerastium maximum. Site 31: alpine basin, 73-46. Weinetal.
(1964) extended the range of C. maximum eastward to the
headwaters of the Yukon River from near the Alaska border
(Hultén 1968). The collection reported here extends the
range further eastward to the Richardson Mountains of
northwestern District of Mackenzie. This species is thus new
to the flora of Continental Northwest Territories (Porsild
and Cody 1968).
Silene acaulis ssp. subacaulescens (Moss Campion). Site 7:
alpine tundra, 74-341; site 30: sphagnum bog, 73-5. Hultén
(1968) plotted a collection locality along the Firth River
drainage near the Alaskan border. The specimens reported
here are the first collections from the predicted range for
Yukon Territory north of 65°N (Hultén 1968).
Ranunculaceae
Anemone narcissiflora Ss. lat. Site 29: sedges on rock slide, 73-
1. Hultén (1968) indicates several collection sites of A.
narcissiflora ssp. interior from the District of Mackenzie
adjacent to the Yukon Territory border. Our collection is the
first report from the predicted range for Yukon Territory
north of 65°N.
Ranunculus Eschscholtzii. Site 25: creek bank, 73-9. Hultén
(1968) reported a collection from District of Mackenzie
adjacent to Yukon Territory. The specimen reported here is
the first collection from Yukon Territory north of 65°N,
extending the known range westward from northwestern
District of Mackenzie to the Barn Mountains.
Papaveraceae
Papaver Hultenii. Site 24: rock outcrop, 75-/. First
collection within the predicted range for northern Yukon
Territory (Hultén 1968).
Papaver Macounii (P. Keelei). Site 7: White Spruce stand,
74-618; site 10: south-facing slope, 74-161; site 27: creek
bank, 70-22; site 30: sphagnum bog, 73-4. Hultén (1968)
indicated collection localities from adjacent areas in Alaska
and northwestern District of Mackenzie. These are the first
specimens collected in the predicted range for Yukon
Territory north of 65°N (Hultén 1968).
NAGY ET AL.: NORTHERN VASCULAR PLANTS
263
Cruciferae
Cardamine umbellata. Site 1: lakeshore, 74-641. First
collection from Yukon Territory north of 65°N, extending
the known range significantly northward to the Barn
Mountains (Hultén 1968).
Draba alpina. Site 7: dry rocky ridge, 74-100. Welsh and
Rigby (1971) reported a collection from the Firth River
drainage, within the range predicted by Hultén (1968). Our
collection extends the known range southeastward from this
station.
Draba borealis. Site 24: rock outcrop, 75-4. Hultén (1968)
plotted an isolated collection in southwestern Yukon
Territory from about 137° W. A more recent map is given by
Mulligan (1970). The specimen reported here is the first
collection from northern Yukon Territory, and extends its
known range significantly northward from about 63°N.
Draba crassifolia. Site 24: rock outcrop, 75-9. Hulten (1968)
reported an isolated collection from the Mackenzie River
drainage adjacent to northern Yukon Territory. A more
recent map is given by Mulligan (1975). The specimen
reported here is the first collection from Yukon Territory
north of 65°N, extending the known range westward.
Draba incerta. Site 7: dry rocky ridge, 74-98; site 10: south-
facing slope, 74-158, 74-164; site 13: dry rocky ridge, 74-404.
Hulten (1968) plotted three collections from southern Yukon
Territory in close proximity to the British Columbia border.
A more extensive map is given by Mulligan (1972). The
collections reported here extend the known range signifi-
cantly northward to northern Yukon Territory.
Droseraceae
Drosera rotundifolia (Round-leaved Sundew). Site 18:
sphagnum bog, 74-543. Hultén (1968) reported an isolated
collection from the Mackenzie River drainage. Our specimen
is the first collection from Yukon Territory north of 65°N,
extending the known range northward to the Old Crow
Flats.
Saxifragaceae
Saxifraga caespitosa (Tufted Saxifrage). Site 11: dry rocky
ridge, 74-183. Extends the known range southeastward from
the recent first collections reported from northern Yukon
Territory by Welsh and Rigby (1971) and Wein et al. (1974).
Saxifraga hieracifolia. Site 21: tundra, 74-587. Hultén (1968)
and Wein et al. (1974) reported collections from localities
along north coastal Yukon Territory. The collection
reported here extends the known range slightly southward
along the Firth River drainage.
Ribes triste (Red Currant). Site 4: alder stand on river bank,
74-190. Previously reported from the Old Crow Flats and
northwestern District of Mackenzie by Hultén (1968). The
specimen reported here extends the known range northward
to the arctic coastal plain.
Rosaceae
Dryas octopetala L. ssp. alaskensis (D. alaskensis). Site 7:
White Spruce stand, 74-82; site 27: dry ridge, 70-22. Wein et
al. (1974) reported the first collection of this species for
northern Yukon Territory from the headwaters of the Yukon
River drainage. The collections reported here extend the
264
known range northward and bridge the gap between
collection localities in Alaska and northwestern District of
Mackenzie (Hultén 1968).
Potentilla elegans. Site 3: dry rocky slope, 74-36; site 6:
alpine tundra, 74-350; site 11: dry rocky ridge, 74-179, site 13:
drainage, 74-377. Extends the known range eastward from
the recent extensions reported by Welsh and Rigby (1971)
and Wein et al. (1974).
Potentilla hyparctica. Site 11: dry rocky ridge, 74-184; site
14: south-facing slope, 74-453. First collections reported for
Yukon Territory north of 65°N, extending the known range
eastward from localities in Alaska (Hultén 1968).
Potentilla Hookeriana ssp. Hookeriana (P. nivea ssp.
Hookeriana). Site 22: rock crevices, 74-671. Hultén (1968)
reported collections of P. Hookeriana from coastal northern
Yukon Territory and the Old Crow Flats. The collection
reported here extends the known distribution southeastward
within the predicted range (Hulten 1968).
Potentilla rubricaulis. Site 3: dry rocky ridge, 74-24,
74-237; site 7: dry rocky ridge, 74-103, 74-114, 74-257,
74-289, alpine tundra, 74-334; site 8: dry rocky slope, 74-70;
site 10: south-facing slope, 74-154, 74-168. First collections
from Yukon Territory, expanding the known range west-
ward from northwestern District of Mackenzie (Hultén
1968).
Rubus arcticus ssp. acaulis (R. acaulis). Site 16: bog,
74-504. Hultén (1968) reported collections from adjacent
areas in Alaska and northwestern District of Mackenzie. The
specimen reported here is the first collection from the
predicted range for Yukon Territory north of 65°N (Hultén
1968).
Sibbaldia procumbens. Site 6: alpine tundra, 74-360. Hultén
(1968) reported an isolated collection from northwestern
District of Mackenzie. The specimen reported here is the first
collection from Yukon Territory north of 65° N, and extends
the known range westward to the Barn Mountains from the
Richardson Mountains in northwestern District of Mac-
kenzie.
Leguminosae
Oxytropis borealis (O. glutinosa). Site 24: rock outcrop,
75-18. First report from the range predicted for northern
Yukon Territory, and helps bridge the gap in its distribution
between localities in northwestern District of Mackenzie and
Alaska (Hultén 1968).
Oxytropis deflexa var. sericea. Site 16: bog, 74-496. First
collection from Yukon Territory north of 65°N, extending
the known range northward to the Old Crow Flats (Hultén
1968).
Oxytropis nigrescens ssp. pygmaea ( O. pygmaea). Site 7: dry
rocky ridge, 74-55, 74-109; site 11: dry rocky slope,
74-174; site 26: dry hilltop, 70-20h; site 29: sedge association
on rock slide, 73-0. Hultén (1968) reported several collec-
tions from the Alaskan coast adjacent to the Yukon
Territory border. The collections reported here extend the
known range across northern Yukon Territory.
Pyrolaceae
Moneses uniflora (One-flowered Pyrola). Site 7: White
THE CANADIAN FIELD-NATURALIST
Vol. 93
Spruce stand, 74-605. Hultén (1968) plotted a collection site
along the Yukon River drainage near the Alaska border. Our
collection extends the known range northward to the Barn
Mountains.
Pyrola secunda ssp. obtusata (One-sided Pyrola). Site 7:
White Spruce stand, 74-617; site 16: bog, 74-493; site 17:
White Spruce stand, 74-525. Hultén (1968) indicated a
collection site from the Firth River drainage along the
Alaska border. The specimens reported here are the first
collections from the predicted range for northern Yukon
Territory.
Ericaceae
Chamaedaphne calyculata (Leather-leaf). Site 19: sphagnum
bog, 74-562. Hultén (1968) plotted a collection site along the
Yukon River drainage near the Alaska border. The
collection reported here extends the known range north-
ward, well onto the Old Crow Flats.
Ledum palustre ssp. groenlandicum (L. groenlandicum)
(Labrador-tea). Site 15: White Spruce stand, 74-474. Hultén
(1968) reported a collection from the Yukon River drainage
near the village of Old Crow. Our collection extends the
known range northward to the Barn Mountains.
Boraginaceae
Eritrichium splendens. Site 7: dry rocky ridge, 74-261,
74-306. Hultén (1968) and Wein et al. (1974) reported
specimens from the Yukon River drainage near the Alaska
and District of Mackenzie borders, respectively. The
collections cited here extend the known range northward to
the Barn Mountains in northern Yukon Territory.
Scrophulariaceae
Pedicularis lapponica. Site 1: tundra, 74-199, 74-234;
site 9: tundra, 74-222; site 26: dry hilltop, 70-20L. Hultén
(1968) reported a collection from the Old Crow Flats in
northern Yukon Territory. The collections reported here
extend the known distribution northward to the Barn
Mountains, but still fall within the predicted range (Hultén
1968).
Lentibulariaceae
Pinguicula villosa. Site 1: polygonal ground, 74-268; site 18:
sphagnum bog, 74-552; site 26: dry ridge, 70-20m. Most
northern collections within the range predicted by Hulten
(1968). He had plotted a collection site on the Yukon River
drainage near the southern edge of the Old Crow Flats.
Utricularia vulgaris (Bladderwort). Site 18: submerged
aquatic, 74-555. First collection from the predicted range for
Yukon Territory north of 65°N (Hulten 1968).
Rubiaceae
Galium Brandegei. Site 19: sphagnum bog, 74-558. First
collection from the predicted range for the Yukon Territory
and is new to the Territory. Helps bridge the gap in the
distribution between collection localities in Alaska and
northwestern District of Mackenzie (Hultén 1968).
Caprifoliaceae
Linnaea borealis ssp. americana (Twinflower). Site 3: dry
rocky slope, 74-630. first collection from its predicted range
for Yukon Territory north of 65°N.
1979
Compositae
Aster alpinus ssp. Vierhapperi. Site 3: dry rocky ridge,
74-283; site 7: dry rocky ridge, 74-260, 74-272, 74-293.
Wein et al. (1974) reported a recent range extension from the
Yukon River drainage in northern Yukon Territory adjacent
to the District of Mackenzie border. The collections reported
here extend the known range northward to the Barn
Mountains.
Antennaria Friesiana ssp. compacta (A. neoalaskana). Site
3: dry rocky ridge, 74-239; site 7: dry rocky ridge,
74-113; site 8: dry rocky ridge, 74-3/1b; site 10: south slope,
74-144; site 13: dry rocky ridge, 74-405, 74-4179. Hultén
(1968) reported a collection from the Old Crow Flats. The
specimens reported here are the northernmost collections
within the range predicted by Hultén (1968) for northern
Yukon Territory.
Senecio atropurpureus ssp. tomentosus (Kjellm.) Hult.
(S. Kjellmannii). Site 11: dry rocky ridge, 74-186. First
known collection from its predicted range in Yukon
Territory north of 65°N, and helps bridge the gap in the
distribution between collection localities in Alaska and
District of Mackenzie.
Taraxacum phymatocarpum. Site 7: rocky alpine tundra,
74-105, 74-325, 74-338. Hultén (1968) reported a collection
from Herschel Island. These collections extend the known
distribution within the predicted range, southward to the
Barn Mountains.
Acknowledgments
The research was undertaken as part of a broader
study of the ecology of the arctic mountain Grizzly
Bear under the auspices of the Canadian Wildlife
Service. We thank Anne Gunn, Canadian Wildlife
Service, who read the manuscript and provided
helpful suggestions.
NAGY ET AL.:. NORTHERN VASCULAR PLANTS
265
Literature Cited
Bostock, H.S. 1961. Physiography and resources of the
northern Yukon. Canadian Geographical Journal 63:
112-119.
Cody, W. J.and A. E. Porsild. 1968. Additions to the flora
of Continental Northwest Territories, Canada. Canadian
Field-Naturalist 82: 263-275.
Hultén, E. 1968. Flora of Alaska and neighboring terri-
tories. Stanford University Press, Stanford, California.
1008 pp.
Mulligan, G. A. 1970. Cytotaxonomic studies of Draba
glabella and its close allies in Canada and Alaska.
Canadian Journal of Botany 48: 1431-1437.
Mulligan, G. A. 1972. Cytotaxomomic studies of Draba
species in Canada and Alaska: D. oligosperma and
D. incerta. Canadian Journal of Botany 50: 1763-1766.
Mulligan, G. A. 1975. Draba crassifolia, D. albertina, D.
nemorosa, and D. stenoloba in Canada and Alaska.
Canadian Journal of Botany 53: 745-751.
Porsild, A. E. 1951. Botany of the southeastern Yukon
adjacent to the Canol Road. National Museum of Canada
Bulletin 121: 1-400.
Porsild, A. E. 1975. Materials for a flora of Central Yukon
Territory. National Museums of Canada, Publications in
Botany, Number 4. 77 pp.
Porsild, A.E. and W.J. Cody. 1968. Checklist of the
vascular plants of Continental Northwest Territories,
Canada. Plant Research Institute, Canada Department
of Agriculture, Ottawa. 102 pp.
Wein, R. W., L. R. Hettinger, A. J. Janz, and W. J. Cody.
1974. Vascular plant range extensions in the northern
Yukon Territory and northwestern Mackenzie District,
Canada. Canadian Field-Naturalist 88: 57-66.
Welsh, S. L. and J. K. Rigby. 1971. Botanical and physio-
graphic reconnaisance of Northern Yukon. Brigham
Young University Science Bulletin, Biological Series,
Volume XIV, Number 2. 64 pp.
Received 21 September 1977
Accepted 9 March 1979
Food and Feeding of the Rainbow Smelt (Osmerus mordax) in
Lake Simcoe, Ontario
HUGH R. MACCRIMMON and ROBERT W. PUGSLEY
Department of Zoology, University of Guelph, Guelph, Ontario NIG 2W1]
MacCrimmon, H. R. and R. W. Pugsley. 1979. Food and feeding of the Rainbow Smelt (Osmerus mordax) in Lake Simcoe,
Ontario. Canadian Field-Naturalist 93(3): 266-271.
Feeding by adult Rainbow Smelt (Osmerus mordax) is selective and varies with fish size, sex, and season. Intensity of feeding
is lowered prior to spawning. Zooplankton is the major food, and we found no evidence of predation on the eggs or young of
sport fishes. Fish were found in the stomachs of 11% of the females and 4.3% of the males and were mainly young-of-the-year
smelt in summer and lake Emerald Shiners ( Notropis atherinoides) in winter and spring. Fish were eaten only by smelt of age
groups I to V (156 to 204 mm in fork length) while crustaceans were the principal food of young-of-the-year smelt.
Key Words: seasonal variation, selection, food preference, diet, predation, food organisms, Osmerus mordax, limiting
factors, body size, feeding behavior.
With the exception of a few lakes in the eastern part
of the province (Dymond 1944), the invasion of
Rainbow Smelt, Osmerus mordax, into Ontario
inland waters is of recent origin. Smelt first appeared
in Lake Simcoe (44°23’N, 79°18’W) in 1962, pre-
sumably as a result of an unrecorded release about
1960: a substantial naturalized smelt population
resulted (MacCrimmon and Skobe 1970).
Because of a decline in Lake Trout (Salvelinus
namaycush) and Whitefish (Coregonus clupeaformis)
fisheries concurrent with the naturalization of smelt in
Lake Simcoe, and because of the smelt’s unique
dentition (McAllister 1963), it is a popular accusation
that the smelt is a serious predator on the eggs and
young of commercial and sport fishes. This assump-
tion is strengthened by the fact that Lake Simcoe
smelt can be readily caught by anglers using hooks
baited with fresh or salted Emerald Shiners, Notropis
atherinoides, especially in the winter ice fishery.
Studies elsewhere, however, have not confirmed that
the smelt is an important predator on fishes (Creaser
1925, 1928: Kendall 1927: Greene 1930: Schneberger
1936: Gordon 1961: Ferguson 1965: Rupp 1968:
Burbidge 1969: Delisle 1969: Lackey 1969: Anderson
and Smith 1971: Selgeby et al. 1978).
We examined the stomach contents of smelt taken
monthly by gill nets set in the lake and of mature
adults collected from a river spawning area. Seasonal
patterns of food and feeding of smelt in Lake Simcoe
are reported, and the importance of predation on
fishes assessed.
Materials and Methods
The area of intensive study included a 54-km2
section of Lake Simcoe north of Sibbald Park,
extending west from Georgina Island to Jackson’s
Point and then north to the 22-m depth contour.
Monthly samples were taken from this area between
August 1970 and August 1971 by means of gill nets set
in the late morning for periods not exceeding | h.
Specimens were chilled and examined in the labora-
tory within 3 h of capture. An additional sample of
336 smelt was collected from the 1971 spawning run in
the nearby Pefferlaw River and frozen for later study.
The data from fish stomachs taken during the
spawning period are dealt with separately from those
of other months.
Stomach contents were examined under a binocu-
lar microscope at 16X using a technique similar to that
of Thompson (1959), Ferguson (1965), and Burbidge
(1969). The volume of partially digested food was
estimated as a percentage of its contribution to the
total stomach food volume. The amounts of the
individual food items in the remaining volume were
expressed as percentage contribution to the total
volume of undigested recognizable food items. The
frequency of occurrence of specific food items is
expressed as a percentage of those stomachs contain-
ing recognizable items. Stomachs containing only
chyme or fluid were considered to be empty.
Results
Examination of the stomachs of 1416 adult smelt
from Lake Simcoe between August 1970 and 1971
found that 76% contained food and, of these,
recognizable items were present in 91.5% (Table 1).
Feeding varied seasonally, both in kind and quantity
of food eaten. All smelt stomachs collected in early
June contained food, as did over 84% of those
collected in July to December. Food intake was
minimal in March and April before the spawning
period but increased during and after spawning
among mature fish captured during the Pefferlaw
River spawning run. Females resumed feeding more
quickly than males after spawning. Stomach fullness
normally fluctuated between 47 and 69% of stomach
266
1979 MACCRIMMON AND PUGSLEY: RAINBOW SMELT FOOD, LAKE SIMCOE
267
TABLE !—Monthly stomach contents of mature Rainbow Smelt showing the number containing food, fullness, and the
occurrences of recognizable food, all expressed in percentages
Stomachs
Stomachs Stomach containing
Sample containing fullness recognizable
Month size food by volume food items
fe) ? 3 2 fe) 2 fe) 2
Aug. 22 89 100.0 98.1 50.9 46.7 90.9 92.1
Sept. i 18 29 100.0 96.6 51.7 55.9 50.0 Sal
Oct. 37 68 100.0 100.0 SPT 50.9 89.2 77.9
Nov. ] 36 — 86.1 — 68.6 — 96.8
Dec. Mp) 78 100.0 98.7 S33 60.0 100.0 98.7
Jan. 72 60 100.0 100.0 57.9 62.6 94.4 98.3
Feb. 48 60 72.9 71.7 36.3 44.9 97.1 88.4
Mar. 54 61 16.7 21.3 55.0 46.5 100.0 100.0
Apr.* 163 62 Die) 38.7 10.3 11.3 100.0 100.0
May 28 68 64.3 82.4 58.1 68.5 100.0 100.0
June 46 58 100.0 ~ 100.0 64.0 67.6 87.0 89.7
July 29 83 86.2 86.8 65.2 48.7 92.0 87.5
Aug. 4] 83 97.6 92.8 70.3 59.0 97.5 96.1
By sex
Number 581 835 38] 695 381 695 349 635
Mean 65.6 83.2 S245) 55.0 91.6 91.4
Combined (6+@)
Number 1416 1076 1076 984
Mean 76.0 54.0 91.5
Mature fish on
Pefferlaw River
spawning grounds 336 42.0 45.9 100.0
*Excludes spawning fish.
capacity, but the food volume in April was only 10%.
Although stomach fullness varied slightly (1.0-16.5%)
between males and females in any one monthly
sampling period, the yearly average was nearly the
same (Table 1).
The importance of each food item varied with
season and both sexes typically ate the most abundant
and/or largest organisms available. In winter
(January to March) copepods (Cyclops sp. and
Diaptomus oregonensis) were the most important
food items eaten by both sexes, followed by clado-
cerans (mainly Bosmina sp.) which became prevalent
in February. Amphipods (Gammarus) and fish (N.
atherinoides, and young-of-the-year O. mordax)
made up 28 and 40% respectively, of the food in
February and March. In April, prior to spawning,
copepods and tendipedids were the most frequent
organisms eaten (Table 2).
In smelt captured during the spawning run in the
Pefferlaw River, fish eggs (O. mordax and Cato-
stomus commersoni) accounted for 55.7% of the food
volume in those stomachs containing food. No
extraneous matter was found in the eggs. Emerald
Shiners were found in 31% of these fish, and made up
30% of the total volume of food consumed in the
spawning and post-spawning period. The remaining
volume of recognizable food items consisted of
unidentified fish larvae (2.8%), Gammarus (0.7%),
and algae (0.7%). Empty stomachs were found in 58%
of fish collected and many of these were enlarged and
contained only clear fluid.
Smelt collected in the lake in May and June had fed
principally on the dipteran larvae of Chaoborus
albipes, Tendipes tentans, Pentaneuris monilis, and
the ephemeropteran nymph of Hexagenia sp. No
bottom detritus was present in stomachs containing
these benthic organisms. In July, cladocerans ( Lepto-
dora kindtii), tendipedidae pupae, and copepods were
the dominant food items. In September to December,
fish (mostly preserved Emerald Shiners used to
prebait Whitefish fishing grounds) and cladocerans
(Bosmina sp. and Daphnia longispina) made up a
greater proportion of the food than did copepods
(Table 2).
Although the seasonal variation in food items eaten
was similar in both male and female smelt, slight
differences in stomach volume and frequency of
occurrence of these items did occur. Male smelt ate
268
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 2—Monthly stomach contents of male and female Rainbow Smelt expressed by percent volume and by frequency (in
parenthesis) of occurrence
Month Cladocera Copepoda Tendipedidae
Aug. 53.6 4.6 32.0
(88.1) (82.2) (47.5)
Sept. 49.6 18.9 13.6
(75.0) (78.6) (42.9)
Oct. 49.1 5.8 6.8
(65.1) (58.1) (12.8)
Nov. 76.0 6.0 0.7
(100.0) (100.0) (6.7)
Dec 92.1 dS —
(100.0) (100.0) —
Jan. 5.4 82.9 0.3
(80.3) (92.1) (3.1)
Feb. 36.0 30.7 0.3
(70.8) (79.2) (1.4)
Mar. 0.5 55.0 —
(4.5) (59.1) =
Apr. — 69.9 14.2
— (71.1) (21.3)
May _ 8.5 46.5
(25.0) (67.7)
June 2.0 19.7 52.0
(12.0) (25.0) (53.3)
July 46.5 9.4 IW 3a
(83.7) (45.3) (43.0)
Aug. 80.3 DP? 11.0
(91.2) (30.1) (31.9)
Mean, sexes
combined 39.5 23.9 15.8
(64.4) (64.0) (26.3)
Mean, by sex
3 36.0 35.9 14.2
(66.2) (70.5) (27.0)
S 41.5 17.3 16.6
(63.5) (60.5) (26.0)
Food item
Fish Chaoborus Hexagenia Gammarus
6.0 3.0 — —
(6.9) (3.0) — —
17.1 _ 0.4 _—
(21.4) — (3.6)
34.1 _ — 3.0
(37.2) = — (5.8)
11.0 — — 6.3
(16.7) — — (16.7)
— — —_— 0.4
— _ _ (1.0)
7.1 0.2 — 0.9
(@A))) (2.4) _ (7.9)
11.0 5.1 _ 16.8
(12.5) (6.9) —_ (19.4)
26.1 3.2 _ 14.1
(36.4) (4.5) — (18.2)
1.7 oo _ 3.3
(1.6) _ — (4.9)
4.4 9.3 24.3 4.3
(4.4) (27.9) (30.9) (5.9)
+ 2.1 28.0 =
aa (6.4) (38.0) —
Dell DAES 6.9 =
(2.3) (46.5) (8.1)
Peo) 3.8 -- —
(2.7) (16.8) — —
7.6 33.9) 49 2.6
(8.6) (9.6) (3.0) (3.6)
315) 2.1 48 2.8
(4.3) (8.0) (1.2) (3.4)
9.9 49 5.0 2.4
(11.0) (10.7) (5.4) (3.9)
cladocerans and copepods in equal quantities (36% by
volume), supplemented by tendipedidae (14.2%) and,
to a lesser extent, by Hexagenia, fish, Gammarus, and
Chaoborus, respectively. Females fed primarily on
Cladocerans (41.5%), followed, in decreasing
amounts, by copepods, tendipedidae, fish, Chao-
borus, Hexagenia, and Gammarus (Table 2).
Larger smelt exhibited a more varied diet than did
smaller fish (Table 3). The occurrence of tendipedidae
(larvae and pupae), Chaoborus, Hexagenia, and fish
(Emerald Shiners and young-of-the-year Rainbow
Smelt) each increased with fish length. Fish first
appeared in the diet of smelt at Age Group |.
Most of the identifiable fish eaten throughout the
year were young-of-the-year smelt. During the
autumn and winter months, preserved (salted)
Emerald Shiners, which are used as bait and for
prebaiting Whitefish fishing grounds, were eaten
occasionally. In the spring (mostly after the smelt had
spawned), live shiners were consumed in substantial
quantities (30.8% by volume in the river sample). The
high percentage of Emerald Shiners occurring in the
October sample (34% by volume) resulted from a
collection of smelt made in proximity to an area newly
baited for Whitefish. The only other fish found
in smelt stomachs were two larval White Suckers (C.
commersoni) (15 mm in length) and a single small
darter (Etheostoma sp.).
Discussion
As a result of the broad temperature regime
favorable for smelt feeding and growth (Ferguson
1965), the smelt is able to feed throughout the year and
at various water depths on a variety of food
1979 MACCRIMMON AND PUGSLEY: RAINBOW SMELT FOOD, LAKE SIMCOE 269
organisms, especially zooplankton supplemented sea-
sonally by small fish and benthic invertebrates.
Similar items of diet are commonly used by smelt in
the Great Lakes (Kendall 1927: Schneberger 1936:
Baldwin 1948: Hale 1960: Gordon 1961; Anderson
and Smith 1971) and in other inland waters (Greene
1930; Reif and Tappa 1966: Rupp 1968; Burbidge
1969: Delisle 1969; Lackey 1969). The diet of
landlocked Arctic Smelt (O. mordax dentex) in the
Rybinsk reservoir in Russia (Inanova et al. 1970) also
resembles that of freshwater Rainbow Smelt in Lake
Simcoe.
A comparison of the stomach contents of male and
female fish indicates that although the food items are
similar, the females eat more of the larger organisms
(fish and dipteran larvae) and havea more varied diet.
Also, recently spent males are slower to resume
feeding than are females. There is no mention of these
phenomena elsewhere in the published literature.
The absence of debris in smelt stomachs that
contain eggs in the spring months, and benthic larvae
and pupae at most times, indicates that feeding is
selective in Lake Simcoe. Adult smelt generally select
the largest organisms available and the larger fish
have a more complex diet than the smaller fish.
Similar feeding behavior was observed among smelt
in lakes Superior (Hale 1960), Huron (Baldwin 1948),
Champlain (Green 1930), and Gull Lake, Michigan
(Burbidge 1969).
Lake Simcoe smelt feed extensively on eggs and fish
during the spawning and post-spawning season in the
rivers. Although over half the volume of food
consumed at those times was comprised of smelt and
sucker eggs, eggs were not found in smelt stomachs at
other times. Creaser (1925) also found small quanti-
ties of smelt eggs in the stomachs of smelt captured in
the spawning run in Cold Creek, Michigan.
Stomachs of 12% of the adult smelt taken on their
spawning grounds in the Pefferlaw River contained
Emerald Shiners. At other times of the year fish were
found in the stomachs of 8.6% of the adult smelt
[EOE SENT examined and were primarily young-of-the-year smelt
in summer and fall, and salted Emerald Shiners in
winter. The selection of fish at certain seasons has
been reported in other papers. For example, 97.3% of
the stomachs of 147 smelt taken in Crystal Lake,
Michigan in September contained Emerald Shiners
(Creaser 1925). Larvae of the Cisco (Coregonus
artedii) were found in 17% of the stomachs of smelt
taken in May from Black Bay of Lake Superior
(Selgeby et al. 1978). In Lake Huron, Emerald Shiners
were found in 8% of smelt stomachs but only in July
and August, and young-of-the-year smelt were found
OTN OR OD os in 23% of smelt stomachs but only in September
ay yall ae el” sale (Gordon 1961). In New York State, the food of the
S22 SSSSefa8 large race of Lake Champlain smelt contained a lower
Gammarus
10.0
4
6.7
Fish
4.]
10.3
14.3
26.7
Hexagenia
|
I
13
50.0
Food item
18.4
19.0
26.7
25.0
Copepoda Tendipedidae Chaoborus
100.0 -
90.0 _-
89.7
65.6 Wee
54.0 26.4
45.2 28.6
33.3 40.0
25.0 50.0
spawning season
Cladocera
100.0
100.0
97.4
92.6
93.1
83.3
80.0
75.0
Stomachs
containing
recognizable
food items
15
4
320
Total
remains
Fish
empty
3
5
3
2
3
16
of
fish
24
9
79
No.
m)
(m
interval
TABLE 3—Percent annual frequency of occurrence of major food items eaten by adult Rainbow Smelt (130 to 209 mm fork length) excluding fish taken during
Length
SS SS ei ia
270
percentage of fish during June and July, while, in the
Saranac Lakes, fewer fish were eaten in summer
(Greene 1930). Similarly, in Lake Erie, fish occurred
in only 2.3% of smelt stomachs in spring and summer,
but increased to 51.2% during the autumn (Price
1963).
There is no evidence from any study. including this
one. that any segment of the smelt population
becomes totally piscivorous. either on a seasonal or
permanent basis: or that smelt predation is an obvious
factor in the suppression of any sympatric population.
The feeding selectivity of the Rainbow Smelt. how-
ever. could exert competitive pressure on specific food
items required at some critical stage in the develop-
ment of other fish species. For example. during the
summer Lake Simcoe smelt eat more Leptodora
kindtii even though smaller cladocerans are more
abundant. Also Reif and Tappa (1966) suggest that
the differential predation of stocked smelt on zoo-
plankton in Harvey’s Lake, Pennsylvania, has result-
ed in the disappearance of L. kindtii and the
replacement of the larger Daphnia pulex by the
smaller D. dubia. It is possible that competition for
copepods between adult smelt and larval lake Cisco
has led, in part. to the decline of the Cisco in the waters
of western Lake Superior (Anderson and Smith 1971).
Thus. in considering possible impacts of an invad-
ing fish species on the welfare of indigenous species.
such as the relationship between the naturalized smelt
and native Lake Trout (Sa/velinus namaycush) and
Whitefish in Lake Simcoe, a knowledge of relative
diet and food preferences of the species at all stages of
life. as well as ecosystem responses. is essential. The
importance of the observed cannibalism on eggs and
young-of-the-year smelt as a self-regulatory popula-
tion control is not known but in view of the increasing
abundance of the species is obviously not a serious
constraint at the ambient stage of naturalization.
There are insufficient data on the food of other fish
species in Lake Simcoe to permit consideration of the
likelihood of important interspecific competition for
food. nor is there evidence of predation by Rainbow
Smelt on either the Lake Trout or Whitefish popula-
tion. The significance on the Lake Simcoe biosystem
of observed variations in the extent and composition
of the diet of the Rainbow Smelt relative to season.
sex. and size of fish is worthy of more profound
investigation.
Acknowledgments
We are indebted to the staff of the Lake Simcoe
Fisheries Assessment Unit of the Ontario Ministry of
Natural Resources for their aid and the use of their
facilities. Financial assistance for the study was
provided by the Canadian National Sportsmen’s
Show and the National Research Council of Canada.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Literature Cited
Anderson, E. D. and L. L. Smith, Jr. 1971. A synoptic
study of food habits of 30 fish species from western Lake
Superior. University of Minnesota Agricultural Experi-
mental Station, Technical Bulletin 279. 199 pp.
Baldwin, N.S. 1948. The American Smelt, Osmerus mor-
dax (Mitchill) of South Bay. Manitoulin Island. Lake
Huron. Transactions of the American Fisheries Society
78: 176-180.
Burbidge, R. G. 1969. Age. growth, length-weight relation-
ship. sex ratio. and food habits of American Smelt.
Osmerus mordax (Mitchill), from Gull Lake. Michigan.
Transactions of the American Fisheries Society 98: 631—
640.
Creaser, C. W. 1925. The establishment of the Atlantic
smelt in upper waters of the Great Lakes. Papers of the
Michigan Academy of Science. Arts. and Letters 5:
405-424.
Creaser, C. W. 1928. The food of yearling smelt from
Michigan. Papers of the Michigan Academy of Science.
Arts, and Letters. 10: 427-431.
Delisle, C. 1969. Ecologie. croissance et comportement de
léperlan du Lac Heney. comté de Gatineau ainsi que la
répartition en eau douce au Québec. Ph.D. thesis.
Department of Biology, University of Ottawa. 189 pp.
Dymond, J. R. 1944. Spread of the smelt (Osmerus mor-
dax) in the Canadian waters of the Great Lakes. Can-
adian Field-Naturalist 58: 12-14.
Ferguson, R. G. 1965. Bathymetric distribution of Ameri-
can Smelt. Osmerus mordax. in Lake Erie. University of
Michigan Great Lakes Research Division Publication.
13: 47-60.
Gordon, W.G. 1961. Food of the American Smelt in
Saginaw Bay. Lake Huron. Transactions of the American
Fisheries Society 90: 439-443.
Greene, C. W. 1930. The smelts of Lake Champlain. Jn
A biological survey of the Champlain watershed. Supple-
ment to the 19th Annual Report. 1929. of the New York
Conservation Department. pp. 105-129.
Hale, J. G. 1960. Some aspects of the life history of the
smelt (Osmerus mordax) in western Lake Superior.
Minnesota Fish and Game Investigations. Fish Series
2: 25-41.
Inanova, M.N., I. Ye. Permitin, and S.N. Polovkova.
1970. Seasonal distribution features of adult landlocked
smelt (snetok). Osmerus eperlanus eperlanus morpha
spirinchus Pallas. in the Volga Reach of Rybinsk
reservoir. Journal of Ichthyology 10: 678-685.
Kendall, W. C. 1927. The smelts. U.S. Bureau of Fisheries.
Bulletin 11: 217-375.
Lackey, R. T. !969. Seasonal depth distribution of land-
locked Atlantic Salmon. Brook Trout. landlocked Ale-
wives. and American Smelt in a small lake. Journal of
the Fisheries Research Board of Canada 27: 1656-1661.
MacCrimmon, H.R. and E. Skobe. 1970. The fisheries of
Lake Simcoe. Fish and Wildlife Branch. Department of
Lands and Forests (Ontario Ministry of Natural Re-
sources). 140 pp.
McAllister, D. E. 1963. A revision of the smelt family.
Osmeridae. Bulletin of the National Museum of Canada
191, Biological Series 71. 53 pp.
1979
Price, J. W. 1963. A study of the food habits of some Lake
Erie fish. Bulletin of the Ohio Biological Survey (New
Series). Volume II (No. 1). 89 pp.
Reif, C. B. and D. W. Tappa. 1966. Selective predation:
smelt and cladocerans in Harvey Lake. Limnology and
Oceanography I 1: 437-438.
Rupp, R.S. 1968. Life history and ecology of the smelt.
Osmerus mordax, in inland waters of Maine. Maine
Department of Inland Fisheries and Game. Fisheries
Research and Management Division. Final Report. 36 pp.
Schneberger, E. 1936. The biology and economic impor-
tance of the smelt in Green Bay. Transactions of the
American Fisheries Society 66: 139-142.
MACCRIMMON AND PUGSLEY: RAINBOW SMELT FOOD, LAKE SIMCOE Ti
Selgeby, J. H.. W.R. MacCallum, and D. V. Swedberg.
1978. Predation by Rainbow Smelt (Osmerus mordax) on
Lake Herring (Coregonus artedii) in western Lake
Superior. Journal of the Fisheries Research Board of
Canada 35: 1457-1463.
Thompson, R.B. 1959. Food of the squawfish. Prycho-
cheilus oregonensis (Richardson). of the lower Columbia
River. U.S. Fish and Wildlife Service. Fisheries Bulletin
60 (158): 43-58.
Received 15 January 1979
Accepted 16 March 1979
Song Pattern of the Cypress Hills Population of
White-crowned Sparrows
M. ROSS LEIN
Department of Biology, University of Calgary, Calgary, Alberta T2N IN4
Lein, M. R. 1979. Song pattern of the Cypress Hills population of White-crowned Sparrows. Canadian Field-Naturalist
93(3): 272-275.
A disjunct population of White-crowned Sparrows (Zonotrichia leucophrys oriantha) in the Cypress Hills of Alberta
possesses a distinctive song pattern which differs markedly from that of birds of the same geographic race in the Rocky
Mountains of western Alberta. This suggests that this population is truly isolated and does not exchange individuals with
populations in the main range of the subspecies.
Key Words: White-crowned Sparrow, Zonotrichia leucophrys oriantha, Cypress Hills, Alberta, song dialects.
The Cypress Hills of southeastern Alberta and
southwestern Saskatchewan support an island of
mesic forest habitat surrounded by drier shortgrass
prairie. The plateau averages about 450 m higher than
the surrounding plains, and this higher elevation
results in lower temperatures and higher precipita-
tion. Consequently, the region shelters disjunct
populations of many plants and animals characteristic
of the coniferous forests of the montane or boreal
regions of Canada (Halladay 1965).
Although it seems probable that many of these
disjunct populations, including conifers (Thompson
and Kuit 1976) and land snails (Russell 1951), are
completely isolated from other conspecific popula-
tions because of the problems of dispersal across long
distances of inhospitable terrain, the situation is less
clear for populations of mobile organisms such as
birds. The Cypress Hills Plateau possesses disjunct
populations of at least 12 passerine species, separated
from other populations in the Rocky Mountains or
boreal forest by gaps of 250-300 km. It is likely that
conifer forests became established on the Cypress
Hills via colonization from migrating belts of con-
iferous forest following the receding Laurentide ice
sheet at the end of the Wisconsin glaciation, and were
isolated by the expansion of the grassland steppe
vegetation, beginning about 10 000 yr ago (Thomp-
son and Kuijt 1976). The populations of montane or
boreal birds may have been isolated in the Cypress
Hills at this time, or they may have colonized the area
somewhat later. A third possibility is that these
populations are not truly isolated, but continuously
exchange individuals with conspecific populations in
other areas.
The disjunct population of White-crowned Spar-
rows (Zonotrichia leucophrys) breeding in the Cy-
press Hills is assigned to Z. /. oriantha, a black-lored
race that also inhabits the Rocky Mountains from
southern Alberta and British Columbia south to
northern New Mexico, and west to south central
Oregon and central eastern California (American
Ornithologists’ Union 1957). Although the Cypress
Hills population differs in mensural and plumage
characters from other populations of oriantha, this
variation is of a clinal nature and does not warrant
taxonomic separation (Banks 1964). Local popula-
tions of White-crowned Sparrows, particularly the
Pacific coast races Z. /. nuttalli and Z. |. pugetensis
(Marler and Tamura 1962: Baptista 1975, 1977), may
be characterized by distinctive song “dialects.”
Whether these dialects are an adaptation to reduce
gene flow between populations, as suggested by
Nottebohm (1969, 1970), is unclear, but they certainly
may act as markers for groups of birds with little
exchange of individuals (Baptista 1975).
During the summer of 1978 I had the opportunity to
record songs of White-crowned Sparrows in the
Cypress Hills. This paper describes the song pattern of
this population and uses this evidence to suggest that
this population is indeed isolated.
Methods
Recordings were made in Cypress Hills Provincial
Park, Alberta, 5-8 June 1978. A minimum of 15
individuals were taped along the eastern and northern
sides of Reesor Lake and in the valley of Battle
Creek. Other birds were heard but not recorded. The
sparrows were singing strongly in clumps of bushes,
mostly hawthorn (Crataegus spp.), roses (Rosa spp.),
and Buckbrush (Symphoricarpos occidentalis), ad-
joining patches of open grassland.
Songs were recorded with a Nagra 4.2 tape recorder
and a Gibson E. P. M. parabolic microphone. Sona-
grams were prepared with a Kay Elemetrics 6061B
Sona-Graph, using the wide-band filter and high-
shape setting.
DTZ
1979
Results and Discussion
Of the 15 males whose songs were recorded, 14
possessed the black lores characteristic of oriantha.
One male at Reesor Lake, however, had white or pale
gray lores, a character found in the northern race of the
species, Z. /. gambelii.
Representative songs of each of the 15 males are
shown in Figure |. With one exception, all are similar.
They consist of a more or less segmented introductory
whistle, a pair of warbled syllables, two buzzes, and a
terminal slow trill. Although there is some variation in
the form of the warbled syllables among birds, this is
minor and all are clearly variants of the same pattern.
Approximately 6-10 other males that were heard but
not recorded possessed songs of this common pattern.
The one exception (Figure |O) had a song in which
the terminal trill was absent and the pair of warbled
syllables was replaced by a four-note segment with a
very different sound quality. The absence of the trill
may not be significant because birds frequently sing
incomplete songs; however, none of the songs heard
from this male over a period of about 20 min had the
trill, and all had the four-note segment. The male with
the unusual song was not the individual with the
gambelii phenotype. The latter bird’s song (Figure I C)
was comparable to those of other males of this
population.
The song pattern of White-crowned Sparrows in
the Cypress Hills is markedly different from the
“typical” song of oriantha in the Rocky Mountains of
southern Alberta, the nearest conspecific population.
Figure 2 shows songs of birds from three localities in
the Rockies, spanning a distance of approximately
200 km from Waterton Lakes National Park in the
south to the Highwood Pass in the north. All have
similar songs, with three whistled notes in place of the
warbled syllables of the Cypress Hills birds, and a
terminal trill consisting of simpler notes than those of
Cypress Hills songs. No-.songs resembling those of
Cypress Hills birds have been recorded in the Rocky
Mountains of Alberta.
The sharp differences between the songs of Cypress
Hills birds and those of birds from the Rocky
Mountains of Alberta suggest that there is little, if
any, exchange of birds between these two oriantha
populations, and that the Cypress Hills White-
crowned Sparrows form a discrete population. Be-
cause juvenile birds probably learn their song prior to
dispersing from their natal area (Marler 1970), any
bird immigrating to the Cypress Hills from another
population would be expected to sing a different song
pattern. The apparent absence of songs resembling
those of montane oriantha suggests that there is little
influx of birds from this area.
Other disjunct populations of White-crowned
Sparrows are found on other montane “islands” in
LEIN: CYPRESS HILLS WHITE-CROWNED SPARROWS
23
Montana (Thompson 1978). The song patterns of
these populations are unknown. But the fact that
White-crowned Sparrows are absent from the Sweet-
grass Hills (100 km WSW of the Cypress Hills and
140 km E of the Rocky Mountains) although suit-
able habitat is available (Thompson 1978), also
suggests a low rate of immigration of birds from the
main range into outlying populations.
The general similarity of songs of birds over a wide
area in the Rocky Mountains (Figure 2) argues that
there is considerable exchange of birds between
populations in this region. Studies of oriantha
populations in California and Colorado (Baker 1975;
Orejuela and Morton 1975), however, have demon-
strated the existence of dialects in the songs of
populations separated by only 16 km. This suggests
that population structure, dispersal, and the develop-
ment of song dialects may show a considerable
amount of variation within this single subspecies.
There are several possible explanations for the
presence of a white-lored bird in the Cypress Hills.
Migrant gambelii have been recorded there (Taverner
1927; Rand 1948), and this individual may represent
the result of past introgression. But birds with white
lores have been collected on several occasions far to
the east of the range of gambelii, or any of the other
white-lored races (Rand 1948), and this may be a
character that is exhibited occasionally in generally
black-lored populations.
The one individual with an aberrant song (Figure
10) may have copied the song of another population
(Baptista 1974) or it may reflect an error in song-
learning (Baptista 1975).
Acknowledgments
I gratefully acknowledge the aid of Valerie A.
Haines in recording songs in the Cypress Hills, and the
valuable comments on the manuscript made by Luis
F. Baptista.
Literature Cited
American Ornithologists’ Union. 1957. Check-list of North
American birds. Fifth edition. Port City Press, Inc.,
Baltimore, Maryland.
Baker, M.C. 1975. Song dialects and genetic differences in
White-crowned Sparrows (Zonotrichia leucophrys). Evo-
lution 29: 226-241.
Banks, R.C. 1964. Geographic variation in the White-
crowned Sparrow Zonotrichia leucophrys. University of
California Publications in Zoology 70: 1-123.
Baptista, L. F. 1974. The effects of songs of wintering
White-crowned Sparrows on song development in seden-
tary populations of the species. Zeitschrift fur Tier-
psychologie 34: 147-171.
Baptista, L. F. 1975. Song dialects and demes in sedentary
populations of the White-crowned Sparrow (Zonotrichia
leucophrys nuttalli). University of California Publications
in Zoology 105: 1-52.
274 THE CANADIAN FIELD-NATURALIST Vol. 93
A aaa MAAK
J
|
\\ \\ ee pe i
FIGURE 1. Songs of male White-crowned Sparrows from Cypress Hills Provincial Park, Alberta. 1C is the song of a white-
lored bird; all others are songs of black-lored birds. Time scale is in seconds. Each division of ordinate scale is 1 kHz.
1979
HAaALe.
Q ama HAAAAAA AS
1.0 2.0
Ss
LEIN: CYPRESS HILLS WHITE-CROWNED SPARROWS
2S
Seg — F
& cae HAAKKaS
cast
44444444
FIGURE 2. Representative songs of oriantha White-crowned Sparrows from the Rocky Mountains of Alberta. 2A and 2B are
from Waterton Lakes National Park; 2C and 2D are from the Sheep River Valley; and 2E and 2F are from Highwood
Pass, Kananaskis Valley. Time scale is in seconds.
Baptista, L. F. 1977. Geographic variation in song and
dialects of the Puget Sound White-crowned Sparrow.
Condor 79: 356-370.
Halladay, I. R. 1965. Recent biota of the Cypress Hills
Plateau: a general survey of the natural history. Jn Cypress
Hills Plateau guidebook, Part 1. Edited by R. C. Zell, 15th
Annual Field Conference, Alberta Society of Petroleum
Geologists. Pp. 37-54.
Marler, P. 1970. A comparative approach to vocal learn-
ing: song development in White-crowned Sparrows.
Journal of Comparative and Physiological Psychology,
Monograph 71: 1-25. :
Marler, P. and M. Tamura. 1962. Song “dialects” in three
populations of White-crowned Sparrows. Condor 64:
368-377.
Nottebohm, F. 1969. The song of the Chingolo, Zono-
trichia capensis, in Argentina: description and evaluation
of a system of dialects. Condor 71: 399-315.
Nottebohm, F. 1970. Ontogeny of bird song. Science 167:
950-956.
Orejuela, J. E. and M.L. Morton. 1975. Song dialects in
several populations of Mountain White-crowned Spar-
rows (Zonotrichia leucophrys oriantha) in the Sierra
Nevada. Condor 77: 145-153.
Rands, A. L. 1948. Birds of southern Alberta. National
Museum of Canada, Bulletin Number 111.
Russell, L.S. 1951. Land snails of the Cypress Hills and
their significance. Canadian Field-Naturalist 65: 174-175.
Taverner, P. A. 1927. Some recent Canadian records. Auk
44: 217-228.
Thompson, L.S. 1978. Species abundance and habitat
relations of an insular montane avifauna. Condor 80::
1-14.
Thompson, L.S. and J. Kuijt. 1976. Montane and sub-
alpine plants of the Sweetgrass Hills, Montana, and their
relation to early postglacial environments of the northern
Great Plains. Canadian Field-Naturalist 90: 432-448.
Received 10 January 1979
Accepted 26 February 1979
Nesting Biology and Development of Young in Ontario Black Terns
ERICA H. DUNN
Department of Biology, Trent University, Peterborough, Ontario
Present address: Long Point Bird Observatory, P.O. Box 160, Port Rowan, Ontario NOE 1MO
E. H. Dunn. 1979. Nesting biology and development of young in Ontario Black Terns. Canadian Field-Naturalist
93(3): 276-281.
Habitat, nesting substrate, egg characteristics, and growth are described for Black Terns ( Chlidonias niger) nesting on Lake
Erie. Black Terns prefer moderate density of emergent vegetation (mainly Typha) in about | m of water, adjacent to open
water, and return to the same nest site or area as long as it is suitable. Egg and growth characteristics are similar to those for
other temperate-nesting tern species. Minnows form a good portion of the biomass consumed by chicks in this area.
Key Words: Black Terns, Chlidonias niger, nesting, growth, egg characteristics.
Black Terns (Chlidonias niger) nest widely through-
out southern Canada, but their breeding biology is not
well known. As the preferred habitat is cattail (Typha
spp.) marsh with a metre or more of water, and the
young after the first 2 or 3 d leave the nest when
disturbed, accessibility for observation is restricted.
Only one intensive study of breeding biology has been
done (Cuthbert 1954), and this included few data on
development of young. This paper fills in some gaps in
knowledge of nesting biology and gives the first data
on growth of chicks.
Methods
This study area was an approximately 15-ha marsh
on the north side of Long Point, near Port Rowan,
Ontario (42°35’N, 80°24’W). A description of the
vegetation and bird life is given in Dunn and Nol
(1976), and mammals seen in the area were Muskrats
(Ondatra zibethicus) and Mink (Mustela vison).
A canoe was used to traverse the study area. There
were approximately 30-40 Black Tern nests in the
area, in small groups. Twenty-three nests in four
groups were studied in 1975 and 32 in five groups in
1976.
Nests were identified with tagged 2-m stakes pushed
into the mud. Eggs were marked with indelible felt-tip
marker, and measured with dial calipers to the nearest
0.01 mm (length and width at broadest point), and
were weighed several times during incubation to the
nearest 0.1 g with a Pesola spring balance. Occasion-
ally eggs were floated ina beaker of fresh lake water to
determine degree of development (Hays and LeCroy
1971).
Fences of hardware cloth were placed around the
nests during incubation to prevent the young from
leaving the nest site during development. The fences
were 0.4—0.5 m in diameter and 0.3 m high. The adults
readily incubated and cared for young within the
fences, and early placement made it easier to catch
adults later in roofed traps of similar size, with
entrance holes in the top. Adults and young were
banded with U.S. Fishand Wildlife Service aluminum
bands. Color bands were used on adults until it
became clear that they were almost never visible.
Chicks were weighed every | or 2 d to the nearest
0.1 g, and tarsus and ulna were measured with the
adjacent bones held at right angles. Measurements of
skull were from bill tip to back of the skull, with the
ruler held parallel to the bill; and of culmen from bill
tip to feather line on the upper mandible. Sheath and
feather length were measured for the 9th primary,
outer rectrix, and a few body feathers on the anterior
parts of dorsal and ventral tracts.
On four occasions, observations of nests were made
from a blind mounted permanently on an anchored
rowboat.
Results and Discussion
Black Tern nests were formed of dead cattail, most
commonly on a pre-existing mat of floating dead
TABLE |—Percentage of Black Tern nests built on various
substrates: A, Ontario (N = 24)—this study; B, Michi-
gan (N = 23)—Cuthbert (1954); C, Iowa (N = 197)—Berg-
man et al.; D, California (N = 40)—Gould (1974)
A B (Cc D
Floating dead vegetation 75 48 11 15
Floating boards or logs 17 22 33
Cattail rootstock 8 53
Muskrat-built structures 22 36 33
Abandoned birds’ nests” 20
Broken-down bulrushes 9
“Muskrat lodges or feeding platforms of freshly cut vegeta-
tion.
*Platforms of old grebe ( Podiceps) and Forster’s Tern nests.
Black Terns have also been found nesting on abandoned
American Coot (Fulica americanus) nests (Cuthbert 1954).
276
1979
cattail lodged in emergent vegetation (Table 1). No
nests were found on Muskrat houses or feeding
platforms, although these were abundant. The
majority of substrates were floating, in water 1-1.2 m
deep. A few nests in a nearby marsh were found on
isolated small patches of soft mud, but these nest sites
were otherwise normal (see below). Black Tern nests
on dry land are rare (Cuthbert 1954; Richardson
1967). These terns are opportunistic in the choice of
supporting structure for their nests, althougn in my
study floating vegetation and boards seemed to be
preferred over Muskrat houses (Table 1). Previous
papers have not commented on the relative avail-
ability of other substrates.
Nest characteristics were described for 25 nests.
Typically, they were about 2-5 cm high and 25 cm
across. The underlying mats of dead cattail were
lodged in living cattail, usually with heavy growth on
at least one side, and averaged 8 m? (ranging from 0
for a nest on a board to 417 m? in area). Open water
was, on average, 4 m away (range: 0.5-12 m). Some
nests were in large pools, and these were generally on
boards or rootstock, depending less on emergent
vegetation to support the nest substrate. Moderate
cattail growth seemed preferred (standing at least | m
above the water but dispersed enough so that a canoe
could be forced through). A few nests were in thin new
growth, but were rare in dense old stands. Other
studies show that the species composition of aquatic
vegetation at Black Tern breeding sites varies
markedly, but that its density 1s usually moderate.
Water is usually | m deep and nests normally are
adjacent to open water (Cuthbert 1954; Richardson
1967; Bergman et al. 1970: Gould 1974).
Although nests were usually well spaced (only 2 or 3
within a 25-m2 area), a few were within 3 m, with a
clear view between them. Groups had up to 10 nests,
and probably resulted from chance distribution of
good sites. About one quarter of all nests were not
near other tern nests. This dispersion of nests is typical
(Cuthbert 1954: Gould 1974), although nests may
occasionally be much closer together (Hoffman 1926).
Because floating vegetation became waterlogged by
late July and winter storms altered nest-site charac-
TABLE 2—Success of Black Terns nests
DUNN: BLACK TERN NESTINGS
27),
teristics, and because water levels and cattail growth
alter over a period of years, terns must seek new nest
sites each year. McNicholl (1975) predicted that under
these circumstances, the birds would continue to usea
nesting area as long as the habitat is suitable, but when
they must move, they would go as a group to a new
site. This is in contrast to the site tenacity demon-
strated by several gull and tern species with permanent
nesting areas.
A few re-trap data suggest that Black Terns do
return to the same general location of a previous
breeding site and will nest there again if possible. Of
five birds trapped in 1976, which had been banded
previously, one was at nearly the same nest site as in
1975 and another was within the same general locality,
probably moving less than 50 m. (Changes in habitat
between years made exact mapping difficult.) Two
birds banded in 1972 were also found in 1976 within a
short distance of the banding site. The fifth bird
moved farther, about 75-100 m, but was still within
the same patch of marsh vegetation as in 1975.
The Black Terns in my study moved in 1977, after
winter storms had so altered the habitat that no
suitable floating platforms were available for nest
substrates. So many Black Terns nest in the Long
Point marshes, however, that it would have been
impractical to trap other areas to determine whether
McNicholl’s (1975) prediction on group adherence
held true.
The outcome of the 55 nests located during the 2 yr
is given in Table 2. The majority of known losses
occurred in the egg stage. In most cases, eggs
disappeared or did not hatch, and a few nests were
destroyed by storms. Undoubtedly, some losses
resulted from my activities (eggs cracked during
trapping of adults, young in retaining fences eaten by
Mink). Nevertheless, Bergman et al. (1970) showed a
very similar success rate (29% of 192 nests) in a much
larger study with no handling of eggs or birds. There
was no correlation between success and nest-site
characteristics either in my study or in that of
Bergman et al. (1970).
There were no significant differences in measure-
ments of eggs according to sequence in the clutch
1975
No. %
Successful’ 9 39
Unsuccessful 9 39
Undetermined 5) DD,
Total 23
anaes otal sali] Opa amen Total
No. YG No. %
6 19 15 27
23 72 32 58
3 9 8 15
32 55
“Young hatched, but later fate may have been unknown. Nests in which all chicks were known to have died were classified as
unsuccessful.
278 THE CANADIAN FIELD-NATURALIST Vol. 93
TABLE 3—Mean +SD (N) weights and measurements of Black Tern eggs
Egg 1 Egg 2 Egg 3 All eggs
Weight (g) 10.63 + 0.96 (8) 11.42 + 0.51 (6) 10.92 + 0.45 (6) 10.95 + 0.76 (20)
Length (mm) 33.96 + 0.98 (19) 34.32 + 1.45 (13) 34.51+1.13 (20) 34.344 1.32 (91)
Width (mm) 24.88 + 0.63 (19) 25.32 + 0.57 (13) 24.73 £0.45 (20) 24.92 + 0.62 (91)
Shape index* 74.04 + 2.67 (15)
74.22 + 2.84 (12) 72.00 + 2.69 (18) 72.69 + 3.72 (74)
“100 X width/length (Collins and LeCroy 1972).
(Table 3), although third eggs were longer and
narrower and had a lower shape index (Collins and
LeCroy 1972). There was a tendency for the first egg to
be smallest and the second to be largest (Table 4). In
contrast, the smallest egg for Common Terns (Sterna
hirundo) is usually the third one. As in Black Terns,
the third egg is usually the most narrow (Gemperle
and Preston 1955; Gochfeld 1977). Fresh egg weight
of Black Terns was 17.3% of adult body weight, a
slightly lower percentage than shown by the larger
Common and Roseate Terns (S. dougallii) (Collins
and LeCroy 1972).
TABLE 4— Percentage of Black Tern nests in which given eggs
in the laying sequence were largest, of middle size, or smallest
in the clutch. N= 10 clutches. Volume calculated as per
Shott and Preston (1975)
Egg | Egg 2 Egg 3
Largest in clutch 10 60 30
Medium sized 30 30 40
Smallest in clutch 60 10 30
Development of embryos could be followed
through flotation of eggs. The pattern for Black Terns
is very similar to that in Common Terns (Hays and
LeCroy 1971), although Black Tern eggs sometimes
float a day earlier (Table 5). Not enough eggs were
floated to give further details, but within a clutch, the
degree of flotation normally indicated the true laying
sequence. Incubation begins after laying of the first
egg and eggs are usually laid a day apart (Haver-
schmidt 1945), so that the eggs are graded in amount
of incubation.
TABLE 5—Flotation and hatching schedule of Black Tern eggs
Days after
laying of egg Development
6-7 Vertical, sits on pointed end
8 Some eggs float
10-11 Last eggs float
19 - 20 Eggs pip
20 — 22 Eggs hatch
Weights of three freshly hatched chicks averaged
7.25 g, and subsequent growth is detailed in Figures
1-3. The average weight for day 0 (day of hatch)
chicks in Figure | includes birds which may have been
hatched as much as 10-20 h before being weighed.
Weight gain in the chicks shows a pattern very similar
to that of other temperate-nesting species. The growth
curves of terns can best be fitted to logistic equations,
and growth constants calculated for various species
are compared in Table 6. Although Black Terns
appear to grow more rapidly than the other terns, they
70
4.37634 34
te
60
50
Dh
fo}
MEAN ADULT
WEIGHT = 63.4g
(N=27)
WEIGHT (GRAMS)
ol
{o)
20
20
AGE (DAYS)
FIGURE |. Weights of Black Tern chicks. Horizontal bar
shows mean, heavy vertical gives SD, and thin
vertical bar indicates range. The numbers indicate
sample size.
1979 DUNN: BLACK TERN NESTINGS 279
TABLE 6—Comparative growth in northern-nesting tern species
Species K* ‘10 — 90° Adult wt., g Source
Black Tern 0.365 12.1 63 This study
Arctic Tern 0.288 isi 104 Ricklefs 1973
Common Tern 0.300 14.6 125 Ricklefs 1973
0.238 18.5 116 LeCroy and Collins 1972
Roseate Tern 0.237 18.5 108 LeCroy and Collins 1972
Sandwich Tern 0.258 237 Ricklefs 1973
17.0
(Sterna sandvicensis) F
“A constant relating to the growth curve, representing rate at which asymptotic weight is attained. Calculated according to
Ricklefs (1967).
*Time in days between achieving 10% and 90% of asymptotic weight (Ricklefs 1967).
are also much smaller as adults. Comparisons of
growth in relation to asymptotic weight between
Black Terns and other semi-precocial gulls, terns, and
skuas (Table 6; Ricklefs 1973, p. 188) suggest that the
Black Tern has a typical growth rate for its size.
Although few chicks were followed through the
entire growth period, some clearly lagged behind in
weight gain, as is also seen in other tern species (e.g.,
Langham 1972). In contrast to what LeCroy and
Collins (1972) found for Common and Roseate Terns,
however, feather growth in laggard Black Tern chicks
was also retarded. The weight loss which occurred
after attainment of asymptotic weight (close to adult
weight) is also common in other terns (LeCroy and
Collins 1972; LeCroy and LeCroy 1974).
Of the weight and linear measures shown in Figures
1 and 2, no single one proved suitable for aging chicks
accurately in the field, although skull length was
helpful for younger chicks and culmen length for older
ones. Considered all together, these average measure-
ments can be used to estimate the age of an unknown
chick within a 3- to 4-d range. The tarsus reaches full
adult length in the chicks well before fledging, but
skull and culmen must both continue to grow after
first flight (Figure 2).
Growth of the body feathers levelled off by the end
of the weight increase period, but both tail and wing
feathers were still growing rapidly. The latter in
particular can be helpful in aging nestlings (Figure 3),
as was also found for Common and Roseate Tern
chicks (LeCroy and Collins 1972).
The egg tooth in 10 chicks dropped off before day 5,
and in one case, on day | or 2. Typically it seems to
disappear on day 3 or 4, making this species similar to
Common, Forster’s, and Arctic Terns (Sterna para-
disaea), but faster in losing the egg tooth than Roseate
or Least Terns (S. albifrons) (McNicholl 1971:
LeCroy and Collins 1972).
On four occasions, observations of nests with single
chicks were made froma blind mounted ona rowboat.
The watches totalled 7 h for achick 5 d old, 2.25 hfora
15-d-old, and 5.25 h for two 19-d-olds in separate nests.
These watches were too few to support general
conclusions on behavior, but certain observations are
worth noting. The single S-d-old chick was fed more
often than the older ones, and was the only one
brooded. It was fed 5.4 times per hour, as opposed to
3.6 per hour for the 15-d chick and 1.7 times per hour
for each of the two 19-d chicks. Brooding took place
for 14% of the observation period. These figures
correspond well with those of Cuthbert (1954), who
made much lengthier observations of chicks less than
9 d old. There is apparently an increase in feeding rate
over the first few days, then a decline after about 10d,
when most weight gain is complete.
Of 602 feedings observed by Cuthbert (1954), 5% of
the items were minnows, 3% dragonflies and, overall,
93.6% were insects. Of the 56 food items seen in this
study, 13% were minnows, and 6% were dragonflies.
The remainder were usually unidentifiable small
objects (therefore probably not minnows) and seemed
to include many grubs and larvae. Although often
described as primarily insectivorous, Black Terns
clearly use fish as a substantial portion of the food fed
to chicks. Given the large size and digestibility of the:
minnows as compared to insects (Dunn 1973), fish
probably provide at least one-third of the protein
required by the growing chicks.
Begging, feeding, and brooding behaviors corres-
ponded closely to those described in detail by
Cuthbert (1954) and Baggerman et al. (1956). One
parent seemed to bring mainly fish to the nest, while
the other brought small items, often in series of closely
spaced feedings.
Most of the chicks’ non-feeding activity consisted of
preening, walking, or resting, and occasionally pick-
ing up and immediately dropping bits of vegetation.
There was a notable change in behavior by 19 d of
age, when the chicks spent a good deal of time
flapping their wings and hopping into the air. Five
chicks disappeared from the retaining fences on day
19, only one stayed longer; earlier departures were
280 THE CANADIAN FIELD-NATURALIST Vol. 93
TARSUS CULMEN
Ae | Sl 8G he
| 4
al eos \ ee
MEAN ADULT MEAN ADULT
TARSUS = 17.3mm CULMEN = 26.5 mm
(N= 27) (N= 29)
LENGTH (mm)
O)
oO 0)
Je
tt
als
3
50 +
an sales
40 cae
30
MEAN ADULT
SKULL = 59.1 mm
20
(N= 29)
O 3 lO I5 0) © 5 lO I5 20
AGE (DAYS)
FIGURE 2. Measurements of Black Tern chicks. Legend as in Figure 1.
1979
100
80
60
LENGTH (mm)
20
AGE (DAYS)
FIGURE 3. Development of feathers in Black Tern chicks
(total length, including sheath). Legend as in Figure 1.
Upper bars indicate length of 9th primary, lower bars
show outer rectrix growth.
believed to be caused by predation or faulty fencing.
The chick that was underweight throughout the
growth period stayed inside the fence the longest.
Delayed fledging of underweight Common Terns was
noted by LeCroy and LeCroy (1974). I originally
thought disappearances of young represented first
flights, but the last occurrence took place while the
nest was under observation. The chick flapped its
wings and barely hopped the fence, then swam away.
Exact age of fledging is undetermined, but must be
close to 19 d. Baggerman et al. (1956) said flight first
occurred 3 wk after hatching, but Cuthbert (1954) felt
it was closer to 25 d after hatching.
Black Terns are very similar to other temperate-
nesting species in both egg and growth characteristics,
even though other species studied to date are
primarily piscivorous and nest along marine shores
rather than in marshes.
Acknowledgments
This study was supported by the Frank M.
Chapman Memorial Fund in 1975, and in 1976 by the
National Research Council of Canada. The Long
Point Bird Observatory provided facilities, a canoe,
and numerous assistants. I particularly thank Erica
Nol for extensive help in field work, and Marshall
DUNN: BLACK TERN NESTINGS
281
Field for aid in designing traps and giving access to
unpublished banding records.
Literature Cited
Baggerman, B., G. P. Baerends, H. S. Heikens, and J. H.
Mook. 1956. Observations on the behaviour of the Black
- Tern (Chlidonias n. niger(L.)), in the breeding area. Ardea
44: 1-71.
Bergman, R.D., P. Swain, and M. W. Weller. 1970. A
comparative study of nesting Forster’s and Black Terns.
Wilson Bulletin 82: 435-444.
Collins, C. T. and M. LeCroy. 1972. Analysis of measure-
ments, weights, and composition of Common and Roseate
Tern eggs. Wilson Bulletin 84: 187-192.
Cuthbert, N. L. 1954. A nesting study of the Black Tern in
Michigan. Auk 71: 36-63.
Dunn, E. H. 1973. Energy allocation of nestling Double-
crested Cormorants. Ph.D thesis, University of Michigan,
Ann Arbor.
Dunn, E. H. and E. Nol. 1976. /m Cattail marsh. Fortieth
breeding bird census. American Birds 31: 83.
Gochfeld, M. 1977. Intraclutch variation: the uniqueness of
the Common Tern’s third egg. Bird-Banding 48: 325-332.
Gemperle, M.E. and F. W. Preston. 1955. Variation of
shape in the eggs of the Common Tern in the clutch-
sequence. Auk 72: 184-198.
Gould, G.I., Jr. 1974. Breeding success of piscivorous
birds at Eagle Lake, California. M.S. thesis, Humboldt
State University, California.
Haverschmidt, F. 1945. De broedduur van de Zwarte Stern
Chlidonias niger (L.). Ardea 33: 237-240. (English
summary.)
Hays, H. and M. LeCroy. 1971. Field criteria for deter-
mining incubation stage in eggs of the Common Tern.
Wilson Bulletin 83: 425-429.
Hoffman, P. W. 1926. Nesting of the Black Tern in
Wisconsin. Auk 43: 86-87.
Langham, N.P. FE. 1972. Chick survival in terns (Sterna
spp.) with particular reference to the Common Tern.
Journal of Animal Ecology 41: 385-395.
LeCroy, M. and C. T. Collins. 1972. Growth and survival
of Roseate and Common Tern chicks. Auk 89: 595-611...
LeCroy, M. and S. LeCroy. 1974. Growth and fledging in
the Common Tern (Sterna hirundo). Bird-Banding 45:
326-340.
MeNicholl, M. K. 1971. The breeding biology and ecology
of Forster’s Tern (Sterna forsteri) at Delta, Manitoba.
M.S. thesis, University of Manitoba, Winnipeg.
MeNicholl, M.K. 1975. Larid site tenacity and group
adherence in relation to habitat. Auk 92: 98-104.
Richardson, F. 1967. Black Tern nest and egg moving
experiments. Murrelet 48: 52-56.
Ricklefs, R. E. 1967. A graphical method of fitting equa-
tions to growth curves. Ecology 48: 978-983.
Ricklefs, R.E. 1973. Patterns of growth in birds. II.
Growth rate and mode of development. Ibis 115: 177-201.
Shott, A. R. and F. W. Preston. 1975. The surface area of
an egg. Condor 77: 103-104.
Received 30 May 1978
Accepted 27 February 1979
Summer, Autumn, and Winter Diets of Elk in Saskatchewan
HUGH M. HUNT
Wildlife Research Division, Department of Tourism and Renewable Resources, 2602 8th Street East, Saskatoon,
Saskatchewan S7H 0V7
Hunt, H. M. 1979. Summer, autumn, and winter diets of Elk in Saskatchewan. Canadian Field-Naturalist 93(3): 282-287.
The composition of the contents of 88 Elk (Cervus elaphus) rumens obtained in summer, autumn, and winter from the boreal
forest region of east-central Saskatchewan demonstrated that woody browse was the preferred forage of Elk. Relatively
constant amounts of forbs (13-17% of rumen contents) and sedges (Carex spp.) and grasses (7—-8%) were present in the rumens
in each season. Cultivated crops made up over one-quarter of the forage found in autumn rumens. Summer rumen
compositions were characterized by similar high proportions of browse in rumens from each of four areas in east-central
Saskatchewan and slight variations from area to area in the minor amounts of sedges and grasses, forbs, and agricultural
crops. Significant differences from area to area in the individual component foods in each broad forage class in the rumens
demonstrated the variable and adaptable nature of the diet of Elk in Saskatchewan.
Key Words: Elk, food, browse, Saskatchewan, Cervus elaphus.
Extensive habitat alteration has occurred in the
boreal forest of Saskatchewan as a result of increased
timber utilization and accelerated land clearing for
agriculture along the forest fringe. Knowledge of the
seasonal composition of the diet of any wildlife species
is basic to understanding its ecology, particularly in
terms of how habitat changes may affect food
quantity and quality. An investigation of the diet of
Elk or Wapiti (Cervus elaphus) in the boreal forest
was undertaken because other food studies did not
appear to be directly applicable to Saskatchewan Elk.
In a review of 48 studies of the foods eaten by Elk in
the western United States, Kufeld (1973) determined
that grasses were the preferred forage of Elk in all
seasons except summer when forb use was high. In the
only published study of Elk diet in habitat closely
comparable to Saskatchewan’s boreal forest, Riding
Mountain National Park in Manitoba, woody browse
was the preferred forage with grass and grass-like
plants abundant in the diet only in spring and early
winter (Blood 1966).
Study Areas and Methods
From 1973 to 1977, contents of 88 Elk rumens were
sampled from the Porcupine Forest, the Squaw
Rapids area, the Fort a la Corne Wildlife Manage-
ment Unit, and the region surrounding Candle Lake in
east-central Saskachewan (Figure 1). The distribution
_ of rumen samples by area and season is givenin Table
1. The Porcupine Forest is predominantly a Trem-
bling Aspen (Populus tremuloides) forest with some
mixedwood (aspen —- White Spruce (Picea glauca))
portions. The Squaw Rapids area consists of aspen
and Balsam Poplar (Populus balsamifera) stands,
mixedwood stands (aspen — Balsam Poplar — White
Spruce, aspen — Jack Pine (Pinus banksiana)) and
softwood stands (Black Spruce (Picea mariana))
282
along the Saskatchewan River, and the sedge (Carex
spp.) meadow - willow (Salix spp.) complexes of the
Cumberland Delta of the Saskatchewan River.
Aspen, aspen-—Jack Pine, and Jack Pine stands
characterize the Fort ala Corne Wildlife Management
Unit whereas the area around Candle Lake is typical
of the mixedwood section of the boreal forest as
described by Rowe (1972) with aspen, White Birch
(Betula papyrifera), White Spruce, Jack Pine, Black
Spruce, and Tamarack (Larix lJaricina) the main tree
species. Nomenclature for the vascular plants follows
Moss (1959) and Budd and Best (1969); most common
names are taken from Budd and Best (1969).
The mean daily temperature in January in east-
central Saskatchewan is about —20°C; mean July
temperature is about 18°C. Annual preciptation
averages about 42 cm, approximtely two-thirds of
which falls as rain from April to October. Yearly
snowfall averages about 130cm and mid- to late
winter snow depths average about 45 cm.
Causes of death of the sample animals were
collisions with automobiles (8) and trains (1), sport-
hunting (53), unregulated hunting (7), winter mor-
tality (1), and a biological collection program (18).
The rumen samples were from 5 male calves, 48 adult
males, 3 female calves, 23 adult females, and 9 animals
TABLE |—Distribution of rumen samples by area and season
Area
Porcupine Squaw Fort a la Candle
Season Forest Rapids Corne Lake
Summer 28 5) 13 4
Autumn 12 — 3} —
Winter 13 l l 8
Total 53 6 17 12
1979
Nipawin
Saskatchewan
Kilometres
0510 20 30 40
A Candle Lake
B Squaw Rapids
C Fort ala Corne
D Porcupine Forest
SASKATCHEWAN
HUNT: ELK DIETS,
23)
Arbortield Hills
3)
Wildcat
Wilderness
Area
Posquia ——
Preecevi/le
FIGURE 1. Approximate limits of Elk range in east-central Saskatchewan from which the Elk rumens were obtained.
of unknown sex and age.
Most rumen samples were stored by freezing as
suggested by Korschgen (1971) but a few were
preserved in 10% formalin. After thawing, a 1.1-L (1-
qt) aliquot of rumen content was washed through a
5.66-mm mesh screen (U.S. Standard Sieve Series
Mesh No. 3'4) following Dirschl’s (1962) recom-
mendation. The material remaining on the screen was
placed on a white enamel tray, covered with water,
and separated into pre-weighed petri dishes using
forceps. Rumen components were identified macro-
scopically to the lowest taxonomic level possible.
Mushrooms were not identified beyond order, and
grasses were identified only to family (Gramineae);
fine material was sorted by forage class as uniden-
tified herbaceous material or unidentified woody
material. Dry weights of the components were
recorded after they had dried for 24-48 h at 70°C.
Data for the identified portions of the samples were
grouped according to region and season regardless of
sex or age class. The grouped data are presented by the
aggregate percentage method of Martin et al. (1946).
Seasonal groupings were made primarily according to
plant phenological developments described by Stew-
art et al. (1977), and correspond approximately to the
periods 15 June - 30 September (summer), | Octo-
ber — 30 November (autumn), and | December —
30 April (winter). Summer represented the period
following new vegetative growth in spring until killing
frosts halted annual growth, and autumn was the non-
green period following summer before snow depths
became a critical factor influencing forage avail-
284
ability and elk mobility. Winter was the period
characterized by snow cover and lower temperatures
although this period did not end with the disap-
pearance of snow but rather with the commencement
of new vegetative growth.
Results and Discussion
Browse was the major component in the diet of the
Elk in east-central Saskatchewan, especially in sum-
mer and winter (Figure 2). When leaves were available
on the woody plants in summer and autumn, Elk
consumed both leaves and twigs; twigs predominated
in winter diets. Grasses and sedges were of minor
importance in each season and forbs were of moderate
importance. Agricultural crops were eaten when
available in the autumn. Blood (1966) reported similar
use of browse, grasses and sedges, and forbs by Elk in
Riding Mountain National Park, Manitoba.
Elk ate a variety of browse species. Aspen was the
most important browse species both in proportion of
rumen contents and frequency of occurrence (Table
2). Willows, the second most frequently occurring
browse species, were common in rumens from each
season. Wild Red Raspberry (Rubus strigosus), rose
(Rosa spp.), and High-bush Cranberry (Viburnum
trilobum) were common forages in summer although
rose was found frequently in rumens from each
season. The 10 browse species occurring most
SUMMER (n=50)
Browse 49.9°%
WINTER (n=23) AUTUMN (n=15)
FIGURE 2. Composition of the rumen contents from Elk in
summer, autumn, and winter in east-central Sas-
katchewan.
THE CANADIAN FIELD-NATURALIST
Vol. 93
frequently in the rumens were aspen (84% occur-
rence), willow (74%), rose (63%), Wild Red Rasp-
berry (45%), High-bush Cranberry (34%), Red-osier
Dogwood (Cornus stolonifera) (30%), Saskatoon
(Amelanchier alnifolia) (15%), Jack Pine (15%),
Choke Cherry (Prunus virginiana) (13%), and Pin
Cherry (Prunus pensylvanica) (13%). The five most
frequently occurring species accounted for 82% of the
dry weight of the browse identified. Blood (1966)
found rose, aspen, willows, and Saskatoon were the
most important browse items in rumens from Riding
Mountain National Park, making up about 87% of
the total browse segregated. Kufeld (1973) listed
Saskatoon, aspen, Choke Cherry, and willow as
highly valuable Elk forages.
The abundance of most browse species in the
rumens varied seasonally. Use of leaves of woody
species was presumably related to their high levels of
crude protein and total digestible nutrients as demon-
strated by Stewart et al. (1977). Rose hips and the fruit
and leaves of High-bush Cranberry were important
foods in late summer. Significant use of fruits by
White-tailed Deer (Odocoileus virginianus) has been
reported by authors such as Healy (1971), Skinner
and Telfer (1974), and Puglisi et al. (1978). Red-osier
Dogwood was consumed in quantity only in autumn,
and Pin Cherry was common only in summer and
autumn rumens. Similar seasonal use of these two
species by Moose (Alces alces) in Minnesota was
reported by Peek et al. (1976). The frequency of
occurrence of rose in the rumen samples decreased in
winter probably because snow covered this low-
growing shrub.
Forbs were eaten mostly in the summer and autumn
but were almost as important in winter. In Elk rumens
from Riding Mountain National Park, Blood (1966)
found forbs comprised a higher proportion in the
summer (26%) than in this study (17%), but only one-
half the level noted in this study in autumn and winter.
The principal forbs in the Saskatchewan rumens were
Wild Vetch (Vicia americana), Cream-colored Vetch-
ling (Lathyrus ochroleucus), Wild Peavine (Lathyrus
venosus), Prickly Sow-thistle (Sonchus asper), Can-
ada Thistle (Cirsium arvense), Fireweed (Epilobium
angustifolium), and horsetail (Equisetum spp.),
whereas the principal forbs in the Riding Mountain
rumens were Wild Peavine, Cream-colored Vetchling,
Dandelion (Taraxacum officinale), Fireweed, and
Coltsfoot (Petasites sagittatus) (Blood 1966). In
recent cutovers in Saskatchewan, proliferation of
invader or pioneer plant species such as Prickly Sow-
thistle and Canada Thistle provided a source of
forage; similar forages were probably not available to
Riding Mountain Elk as logging is not permitted
there. Blood (1966) concluded that certain forbs in
Riding Mountain National Park were eaten in excess
1979
HUNT: ELK DIETS, SASKATCHEWAN 285
TABLE 2—Detailed seasonal composition of rumen contents from Elk in east-central Saskatchewan. Forages comprising
<1% of the rumen contents are grouped into the “others” categories and the mean proportion of unidentified material in the
samples is also included
Summer (n=50)
Autumn (n=15) Winter (n=23)
Food % Dry % Dry % Dry
categories weight Frequency weight Frequency weight Frequency
Trembling Aspen (Populus tremuloides) 11.9 72.0 20.7 100.0 26.4 100.0
Willow (Salix spp.) ES 70.0 1.5 73.3 15.9 82.6
Raspberry (Rubus strigosus) 11.6 62.0 _ — - 34.8
Rose (Rosa spp.) 12.3 80.0 — 60.0 — =
High-bush Cranberry (Viburnum trilobum) 16.3 48.0 1.7 4.0 — =
Red-osier Dogwood (Cornus stolonifera) o — 8.3 66.7 2 39.1
Pin Cherry (Prunus pensylvanica) 4.4 18.0 Del 13.3 — =
Saskatoon (Amelanchier alnifolia) — — od} 20.0 — —
Jack Pine (Pinus banksiana) 1.3 16.0 1.8 13.3 1.9 13.0
Choke Cherry (Prunus virginiana) a= _ — — 2.4 Dllel/
Honeysuckle (Lonicera spp.) = == = = 1.9 13.0
Others 3.4 _ 3.4 = 2.0 —
Unidentified browse 5.9 44.0 9.1 U3) 19.1 Slee.
Total browse 74.6 49.9 72.8
Sedge (Carex spp.) 2.0 46.0 — 40.0 4.7 39.1
Grass 4.9 68.0 5.8 60.0 82 39.1
Prickly Sow-thistle (Sonchus asper) 1.2 22.0 5.9 33.3 Dey 17.4
Vetchling, Peavine (Lathyrus spp.) 4.2 22.0 — = 1S 13.0
Vetch (Vicia americana) 30) 52.0 — 26.7 = =
Mushroom EY 18.0 — — — —
Fireweed (Epilobium angustifolium) 1.9 36.0 — _— oe
Horsetail (Equisetum spp.) _ — 2.8 13.3 2.2 8.7
Canada Thistle (Cirsium arvense) -- -- — — 4.6 13.0
Alfalfa (Medicago sativa) 19) 2.0 1.6 33) 4.0 4.3
Barley (Hordeum vulgare) — 10.8 40.0 _—
Oats (Avena sativa) a _ 5.9 33.3 _
Wheat (Triticum aestivum) — _ 3.1 6.7 2.0 4.3
Cereal stems — _— 6.0 26.7 —
Others Del — Spl _ DED os
Unidentified herbs _— _ Bal 20.0 _— —
Total herbs 25.4 50.1 DY)
Unidentified material 37.6 96.0 40.4 9353 Dei 86.9
of their availability while some other abundant forbs
made up a small or negligible part of the diet. The
same was probably true for east-central Saskatche-
wan.
Cultivated crops such as Wheat (Triticum aesti-
vum), Oats (Avena sativa), Barley (Hordeum vul-
gare), and Alfalfa (Medicago sativa) were used
extensively in autumn. This may cause significant
conflicts along the forest-farmland interface, par-
ticularly in years of weather-delayed harvests. Elk use
of agricultural crops was not mentioned by Blood
(1966), and Alfalfa is the only agricultural crop
present in the Saskatchewan rumens included in
Kufeld’s (1973) list of Elk forages. Agricultural crops
may be particularly important to Elk in autumn for
maintenance of good body condition prior to the
lengthy period of negative energy balance that
Stewart et al. (1977) describe as characteristic for
Moose in winter.
Mid- to late summer rumen compositions for the
four areas consisted predominantly of woody plant
material (Figure 3). The minor amounts of sedges and
grasses, forbs, and agricultural crops varied slightly
from area to area. Examination of a more detailed
breakdown of the rumen compositions for the four
areas (Table 3) indicated that although the propor-
tions of the browse forage class for each area were
relatively similar, the individual component foods of
the browse and other forage classes differed from area
to area. For example, the proportion of aspen in the
rumens ranged from over half at Candle Lake to only
trace amounts at Squaw Rapids. Willows were
286
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 3—Composition of the rumen contents from Elk in summer in four areas of east-central Saskatchewan. Forages
comprising <1% of the rumen contents are grouped into the “others” categories and the mean proportion of unidentified
material in the samples is also included
Porcupine Squaw Rapids Fort a la Corne Candle Lake
(n=28) (n=5) (n=13) (n=4)
Food % Dry % Dry % Dry % Dry
categories weight Frequency weight Frequency weight Frequency weight Frequency
Trembling Aspen (Populus tremuloides) V3 71.4 = = 13.3 84.6 53.7 100.0
Willow (Salix spp.) 2.9 SA BoP) 60.0 Sal 9253 DES 100.0
Rose (Rosa spp.) 14.5 85.7 11.2 80.0 11.6 76.9 — 50.0
Raspberry (Rubus strigosus) 10.7 53.6 27.6 100.0 10.6 76.9 1.4 50.0
High-bush Cranberry (Viburnum trilobum) — 21.2 60.7 41.4 100.0 1.1 15.4 — =
Balsam Poplar (Populus balsamifera) 2 17.9 = — — — — =
Pin Cherry (Prunus pensylvanica) 3.1 el — — 10.4 53.9 — —
Manitoba Maple (Acer negundo) = 3.6 — —_ — _ — —
Choke Cherry (Prunus virginiana) — — — _ 1.1 15.4 — —
White Birch (Betula papvrifera) 1.0 3.6 — —_— — — — _
Currant, Gooseberry ( Ribes spp.) = = = = = — DT 25.0
Jack Pine (Pinus banksiana) — — — _ 4.2 53.9 2.4 25.0
Others 2.1 ~ 0.4 — 1.1 — 1.0 =
Unidentified browse US 53.6 10.7 80.0 2.6 23.1 — —_
Total browse ales) 9355 Tales 83.1
Sedge (Carex spp.) Dod) 53.6 1.5 60.0 — — 2.4 50.0
Grass 8.0 75.0 — _ 1.5 76.9 — —
Prickly Sow-thistle (Sonchus asper) 1.4 14.3 2.2 20.0 a _ Bo) 75.0
Vetchling, Peavine (Lathyrus spp.) 6.0 32.1 _ 333 Hell - =
Vetch (Vicia americana) 5.3) 57.1 = — 9.2 53.9 — 25.0
Sarsaparilla (Aralia nudicaulis) _ 1.1 20.0 — — = 25.0
Fireweed (Epilobium angustifolium) 3.0 42.9 -— — — 1.3 50.0
Cow-parsnip (Heracleum lanatum) — — — — — = 12 25.0
Mushroom — _ i _ od) 46.2 4.2 25.0
Lichen (Cladonia spp.) — — — — — — 2.9 25.0
Barley (Hordeum vulgare) — _ — — 7.4 Ali — —
Others 0.9 _ 1.7 — Dp? _ Dell _
Unidentified herbs 1.4 21.4 — — — = —
Total herbs 28.5 6.5 28.9 16.9
Unidentified material 41.2 96.4 Dia 100.0 38.6 100.0 22.4 75.0
common in the rumens from the Candle Lake area
and Fort a la Corne but less abundant in the rumens
from the Porcupine Forest and the Squaw Rapids
region. Pin Cherry was a significant component only
in the samples from the Porcupine Forest and Fort a
la Corne; Jack Pine was found only in rumens from
Fort a la Corne and the Candle Lake area. High-bush
Cranberry formed a major part of the food in rumens
from the Porcupine Forest and Squaw Rapids area
but was less common in rumens from the other areas.
Mushrooms formed a large part of the non-woody
forage in samples from both Fort a la Corne and the
Candle Lake area whereas lichens (Cladonia spp.)
appeared in notable quantity only in the rumens from
the Candle Lake region. Deer have also been noted to
eat mushrooms, but the significance of this is
unknown (Healy 1971; Skinner and Telfer 1974). The
cosmopolitan nature of the diet of Elk likely is a factor
that enables them to thrive in many types of habitat.
This study demonstrates the importance of twigs
and leaves of woody plants in the summer, autumn,
and winter diet of Elk in east-central Saskatchewan in
contrast to Kufeld’s (1973) synthesis of Elk diets in the
western United States in which grasses are the
preferred forage. Land clearing along the forest fringe
and subsequent planting of exotic grasses to enhance
cattle production will make those areas less attractive
to Elk as supplies of many of the preferred forages will
be reduced. Clearcut logging in the commercial forest
will stimulate production of most of the preferred Elk
forages but forest managers must ensure that the
proper balance between food and cover is maintained.
1979
PORCUPINE (n=28) SQUAW RAPIDS (n=5)
Sedge,Grass 2.4%
FORT A LA CORNE (n=13) CANDLE LAKE (n=4)
FIGURE 3. Composition of the rumen contents from Elk in
summer from four areas in east-central Saskatch-
ewan.
Acknowledgments
This project was part of a study of the ecology of
Elk in Saskatchewan conducted by the Wildlife
Research Division of the Saskatchewan Department
of Tourism and Renewable Resources. I am indebted
to R. Tennent, D. Arneson, D. Phillips, and B.
Weichel for analysis of the rumen samples. The
assistance of many department members in collecting
rumen samples and for review of the manuscript is
also appreciated.
HUNT: ELK DIETS, SASKATCHEWAN
287
Literature Cited
Blood, D. A. 1966. Range relationships of Elk and cattle in
Riding Mountain National Park, Manitoba. Canadian
Wildlife Service Wildlife Management Bulletin, Series 1,
Number 19. 62 pp.
Budd, A.C. and K.F. Best. 1969. Wild plants of the
Canadian prairies. Canada Department of Agriculture
Publication 983. 519 pp.
Dirschl, H. J. 1962. Sieve mesh size related to analysis of
antelope rumen contents. Journal of Wildlife Manage-
ment 26(3): 327-328.
Healy, W. M. 1971. Forage preferences of tame deer in a
northwest Pennsylvania clear-cutting. Journal of Wildlife
Management 35(4): 717-723.
Korschgen, L. J. 1971. Procedures for food-habits analysis.
In Wildlife management techniques. Edited by R.H.
Giles, Jr. Wildlife Society, Washington, D.C. pp. 233-250.
Kufeld, R. C. 1973. Foods eaten by the Rocky Mountain
Elk. Journal of Range Management 26(2): 106-113.
Martin, A. C., R. H. Gensch, and C. P. Brown. 1946. Al-
ternative methods in upland gamebird food analysis.
Journal of Wildlife Management 10(1): 8-12.
Moss, E. H. 1959. Flora of Alberta. University of Toronto
Press, Toronto, Ontario. 546 pp.
Peek, J. M., D. L. Urich, and R. J. Mackie. 1976. Moose
habitat selection and relationships to forest management
in northeastern Minnesota. Wildlife Monographs, Num-
ber 48. 65 pp.
Puglisi, M. J.,S. A. Liscinsky, and R. F. Harlow. 1978. An
improved methodology of rumen content analysis for
White-tailed Deer. Journal of Wildlife Management 42(2):
397-403.
Rowe, J.S. 1972. Forest regions of Canada. Canada
Department of Environment, Canadian Forestry Service
Publication Number 1300. 172 pp.
Skinner, W.R. and E.S. Telfer. 1974. Spring, summer,
and fall foods of deer in New Brunswick. Journal of
Wildlife Management 38(2): 210-214.
Stewart, R. R.,R. R. MacLennan, and J. D. Kinnear. 1977.
The relationship of plant phenology to Moose. Saskatche-
wan Department of Tourism and Renewable Resources
Technical Bulletin Number 3. 20 pp.
Received 4 November 1978
Accepted 28 March 1979
Taxonomy, Distribution, and Ecology of the Cliff-brake Ferns
(Pellaea: Polypodiaceae) in Alberta
DANIEL F. BRUNTON
Box 59, Canmore, Alberta TOL O0MO
Brunton, Daniel F. 1979. Taxonomy, distribution, and ecology of the cliff-brake ferns (Pellaea: Polypodiaceae) in Alberta.
Canadian Field-Naturalist 93(3): 288-295.
The genus Pellaea in Alberta consists of three species of calcicolous rock ferns. Pellaea atropurpurea, an apogamous triploid,
is rare in the province. It is locally abundant in the Bow Valley of western Alberta, preferring hot arid sites on partially-shaded,
heavily fractured limestone cliff-faces with a southwestern exposure. Pellaea occidentalis, a sexual diploid, is uncommon in
Alberta but is locally abundant in the Bow Valley of western Alberta. It is found on arid south- or southwest-facing, open
limestone cliff-faces. Pellaea suksdorfiana, an apogamous tetraploid, is uncommon in western Alberta where it ranges
throughout the mountains. It prefers cool shaded, east- to north-facing limestone or calcareous sandstone cliff-faces, usually
near water. Pellaea occidentalis and P. suksdorfiana are distinct from the eastern P. glabella, an apogamous tetraploid, on the
basis of distinctive characteristics of range, morphology, cytology, and ecology. Characteristics for separating P.
suksdorfiana, P. occidentalis, and P. atropurpurea are provided. Forty-five previously undocumented records for Alberta
Pellaea are listed.
Key Words: Pellaea atropurpurea, Pellaea occidentalis, Pellaea suksdorfiana, taxonomy, distribution, ecology, cytology,
Alberta, Polypodiaceae.
The cliff-brake ferns ( Pe//aea) have been described
as “.. comparatively rare in Canada or at most locally
abundant...” (Rigby and Britton 1970). This state-
ment still applies to the status of these small calci-
colous rock ferns despite substantial additional study
in parts of Ontario and Quebec (Lafontaine and
Brunton 1972; Brunton and Lafontaine 1974). In
western Canada the genus is represented by three
species, P. atropurpurea(L.) Link, P. occidentalis (E.
Nels.) Butters, and P. suksdorfiana Butters. The latter
two are frequently treated as varieties of the eastern P.
glabella.
This study was initiated to determine whether
ecological evidence exists to support specific status for
the three Alberta taxa of Pe/laea and to gain a better
understanding of its distribution and abundance in
the province.
Taxonomy
The taxa of Pe/laea in Alberta have been described
under several names and/or combinations at a species
and varietal level. Close morphological similarity has
encouraged this, even though there are clear cytolo-
gical differences (Tryon 1957). The three Alberta
species of Pellaea are as follows:
1) Pellaea atropurpurea (L.) Link
2) Pellaea occidentalis (E. Nels.) Butters
= P. glabella Mett. ex Kuhn var. occiden-
talis (E. Nels.) Butters
P. glabella Mett. ex Kuhn var. nana
(Richards.) Cody
= P. pumila Rydb.
3) P. suksdorfiana Butters
= P. glabella Mett. ex Kuhn var. simplex
(E. Nels.) Butters
= P. atropurpurea (L.) Link var. simplex
(Butters) Morton
Few authors today question the validity of P.
atropurpurea as a distinct species (Tryon 1957, and
others). It is an apogamous triploid (n and 2n = 87). It
is also the largest species of Canadian Pellaea. It
occurs across the country from British Columbia to
Quebec and exhibits a distinctive morphology. The
questions arise with P. occidentalis and P. suksdor-
fiana, which are often treated as varieties of P.
glabella.
Pellaea occidentaiis is a sexual diploid (n = 29,
2n = 58) and is a tiny plant of the dry interior of North
America. Pellaea suksdorfiana is an apogamous
tetraploid (n and 2n = 116) which is a species of the
Cordilleran region of western North America. Both
exhibit distinctive morphologies and ecology. The
species with which they are often combined, P.
glabella, is an apogamous tetraploid (n and 2n = 116).
Although maintaining an equivalent cytology to that
of P. suksdorfiana, it is disjunct from that taxon by
over 1300 km. It is quite clearly different in morpho-
logy and cytology from P. occidentalis. Without the
benefit of known ecological differences and with
limited range data available, it is understandable that
many authorities have preferred to treat these as one
species. A. F. Tryon (Gray Herbarium, Harvard
University, personal communication 1977) is in-
288
1979
clined to retain this status for the utility of indicating
relationships in the genus. On the basis of more
complete range data and the demonstration of
ecological preferences of each species which were
observed during our study, we feel that there is
sufficient evidence to justify each taxa being treated
specifically.
’ Banff.
Bt AS :
Mt Mt i
Cory Norquay\
A A
Jasper
National
Park
C ) Unpublished and/or new stations
A Previously published records
National
(Rigby and Britton. 1970)
Park — ~s
kilometres
Kananaskis
Provincial
A
Park ——
BRUNTON: PELLAEA IN ALBERTA
‘Three
A Sisters
289
Distribution and Ecology
Pellaea atropurpurea (Purple Cliff-brake) (Figures 1,
2),
This fern is known sparingly in Canada from British
Columbia to Quebec. It is considered to be the rarest
Pellaea in Canada (Rigby and Britton 1970), although
recently it has been found to be locally abundant in
4
kilometres
” Ga] CALGARY
ye ~~ Bow X ie
angina ne
©
n
eT haa hbridge
Ue
fil ¢
FIGURE |. Distribution of Pe//laea atropurpurea in Alberta.
290
parts of Ontario and western Quebec (White 1977).
Only recently noted in the province by the revision
of a specimen collected in 1900 at Banff (Rigby and
Britton 1970), it is now known from I1 sites in
Alberta. These are all in or near the Bow Valley, alow
dry valley in the eastern slopes of the Rocky
Mountains. The additional stations are listed below.
Herbarium acronyms follow Holmgren and Keuken
(1974) with the exception of DFB which represents the
author’s personal herbarium.
FIGURE 2. Robust clump of Pellaea atropupurea growing in
wide fissure in southwest-facing limestone cliff-face.
Gap Lake, Exshaw, Alberta, 18 February 1978.
4 mi [6.4 km] E of Canmore, Porsild 18147 in
1951 (CAN), Brunton 1289 in 1976 (DFB,DAO,
CAN); 1.5 mi [2.4 km] SE of Exshaw, Bird 18500 in
1967 (UAC), Brunton 1307 in 1977 (DFB); 4 mi
[6.4 km] SE of summit of Mount Head, Bird 28833 in
1972 (UAC); 2.5 km NE of Exshaw, Brunton and
Brunton 1247 in 1976 (DFB,DAO,ALTA,CAN);
2 km NE of Exshaw, Brunton and Grant 1522 in 1977
(DFB,CAN); 2.5 km W of Exshaw, Brunton 1530 in
1978 (DFB); 4.5km W of Exshaw at Gap Lake,
THE CANADIAN FIELD-NATURALIST
Vol. 93
Brunton 1528 in 1978 (DFB,DAO): 1 km NE of
Exshaw, Brunton and Coneybeare 1532 in 1978
(DFB,CAN); 2.5 km S of Kananaskis Environmental
Sciences Center, Barrier Lake, Brunton 1538 in 1978
(DFB); 12 mi[19.3 km] S of Seebe, Cody, Jaques and
Corbin 23037 in 1977 (DAO): southeast Barrier Lake,
Kananaskis Valley, Brunton 1623 in 1979 (DAO).
The species is exceptionally abundant at several
sites, particularly the Grotto Mountain and Gap Lake
sites (Porsild 18147 and Brunton 1289, and Brunton
1528 respectively). Alberta specimens are typically
less than 20 cm tall. Only at the above sites (and at
Barrier Lake, Cody et al. 23037) were plants noted
which approached 30 cm in height, a condition more
typical of eastern Canadian plants. This is likely a
reflection of the much drier climate of Alberta. We
observed that P. atropurpurea in drier-than-normal
sites in Quebec were smaller than the typical.
Alberta P. atropurpurea is found on dry steep
exposed, heavily-fractured limestone rock slopes or
cliffs. With one exception (Heart Mountain, Bird
18500), the stations exhibited a southwestern ex-
posure. Individual plants were partially shaded by
rock ledges and/or scattered woody vegetation,
usually Ribes oxycanthoides and/or Potentilla fruti-
cosa. The plants grow out from under a sheltering
ledge or boulder and are found only in those crevices
in which a considerable amount of soil and/or plant
debris has accumulated. The site preference was very
consistent with other Canadian site descriptions
(Brunton and Lafontaine 1974). The maximum
elevation noted for any station of P. atropurpurea was
1650 m above sea level.
In almost every case, the Woolly Lip-fern (Chei-
lanthes feei) was a close (and frequently abundant)
associate. These warmer-than-normal sites were also
typified by a number of montane species which are
very local in distribution in Alberta. They include
Douglas Fir (Pseudotsuga menzieii), Limber Pine
(Pinus flexilis), and Rocky Mountain Juniper (Juni-
perus scopulorum — possibly J. scopulorum X Juni-
perus Ssp.).
Purple Cliff-brake, being the largest and easiest to
collect of Alberta Pe//aea, is probably represented in
herbaria in greater than natural frequency vis. other
Pellaea.
Pellaea suksdorfiana (Smooth Cliff-brake) (Figures 3,
4).
This species occurs in Canada only in the Cordil-
leran region of British Columbia and Alberta, where it
is considered to be uncommon and local (Taylor 1970:
Rigby and Britton 1970). Prior to this study, it was
known from five locations in Alberta. Twenty
additional stations are listed below:
L979
Spray Falls, Banff N.P., Warren in 1934 (ALTA):
3 mi [4.8 km] W of the Kananaskis Forest Experi-
ment Station, Porsild and Lid 19424 in 1956 (CAN):
5 mi [8.1 km] S of Kananaskis Forest Experiment
Station, Porsild and Lid 19424 in 1956 (CAN); E of
Corral Creek, De Smet Range, Jasper N.P., Porsild
21285 in 1959 (CAN); Talbot Lake, Jasper N.P.,
Porsild 22377 in 1960 (CAN); Bearspaw Dam,
Calgary, A. Birdin 1965 (UAC), Brunton 1534 in 1978
(DFB); entrance near Hinton, Pegg 2095 in 1965
(CAN,ALTA); southwest side Lac des Arc, Bow
BRUNTON: PELLAEA IN ALBERTA
291
Valley, Brunton 1139 in 1976 (DFB); W of Barrier
Pass, Jaques 5059 in 1974 (UAC), Brunton 1293 in
1976 (DAO,DFB): E of Jasper, Jasper N.P., Ogilvie in
1960 (UAC); west side Heart Mountain 1.5 mi
[2.4 km] SE of Exshaw, Brunton 1308 in 1977 (DFB):
2.5 km W of Exshaw, Brunton and Paton 1337, 1338
in 1977 (DFB), Brunton 1531 in 1978 (DFB); 4.5 km
W of Exshaw at Gap Lake, Brunton 1529 in 1978
(DFB); | km NE of Exshaw, Brunton and Coney-
beare 1533 in 1978 (DFB); 12 mi[19.3 km] of Seebe,
Cody, Jaques and Corbin 23038 in 1976 (DAO).
The
‘Three
A Sisters
Jasper
National
Park
@ Unpublished and/or new stations
Banff
National
Park —
A Previously published records
(Rigby and Britton. 1970)
100
kilometres
Kananaskis
Provincial
Park
47 CALGARY
5; a Bow
AD
UF Prato
SPE
FIGURE 3. Distribution of Pellaea suksdorfiana in Alberta.
292
Sight Records: 6.5 km E of Canmore on Grotto
Mountain (Brunton 1977); north side Banff Airfield,
Banff N.P. (Brunton 1978); south side Tunnel
Mountain, Banff N.P. (Brunton 1978); west side Hwy
40 at Barrier Lake (Brunton 1978); east side Hwy 40 at
Barrier Lake (Brunton 1978).
Virtually all of these stations are from cliffs of
limestone. All but two of the stations near Exshaw
(Brunton 1529 and Brunton and Coneybeare 1533)
and one by Barrier Lake (Jacques 5059, Brunton
1293) are adjacent to water bodies. These latter
stations are on north- or northeast-facing slopes.
FIGURE 4. Typical frond of Pellaea suksdorfiana growing out
from fissure in southeast-facing limestone cliff-face.
Gap Lake, Exshaw, Alberta, 18 February 1978.
Pellaea suksdorfiana prefers cool heavily-shaded,
less arid sites than the other Alberta Pellaeas,
normally selecting waterside sites with an eastern
exposure. It often grows in thin fissures in open rock.
Rarely exceeding I5cm in height (and usually
smaller) it is, nonetheless, typically larger than P.
occidentalis. In those sites which are south-facing
(e.g., Brunton and Paton 1338) there is heavy shade
from surrounding trees. Individual clumps are open
and sprawling, with fronds extending well out from
the rock-face.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Pellaea suksdorfiana is uncommon throughout the
Rocky Mountains of Alberta. It extends out onto the
prairie in two sites (near Cochrane and Calgary).
For a morphological comparison of this species
with P. occidentalis see Table 1.
Pellaea occidentalis (E. Nels.) Butters (Western Cliff-
brake) (Figures 5, 6).
This is a species of the dry interior plains and low
mountains of central North America. It is not found
W of Alberta or E of Manitoba in Canada (Rigby and
Britton 1970; Taylor 1970; Tryon 1957). Only in
Wyoming does it extend across the Continental
Divide into areas of moister climate (Hitchcock et al.
1969). It has the most exclusive range of any Canadian
Pellaea, overlapping with others only in southwestern
Alberta. Prior to this study it was known from three
locations in Alberta. Fourteen additional stations are
listed below:
Southwest slope Sawback Range, Banff N.P., Por-
sild and Breitung 15482 in 1946 (CAN); southern
slope Mount Cory, Banff N.P., Porsild 17993 in 1951
(CAN); Kootenay Plains W of Nordegg, Dumais in
1976 (ALTA,DFB); Grotto Mountain, 6.5 km E of
Canmore, Brunton 1290 in 1976 (DFB,DAO); south-
east end Mount Rundle, Canmore, Brunton and
Paton 1328 in 1977 (DFB,DAO): north side Barrier
Lake, Cody, Jaques and Cordin 23043 in 1976 (DAO):
north side Crowsnest Lake, Crowsnest Pass, Brunton
1613 in 1978 (DFB,DAO).
Sight Records: 2.5 km NE of Exshaw (Brunton
1976); 2 km NE of Exshaw (Brunton 1977); 4.5 km W
of Exshaw at Gap Lake (Brunton 1978); south side
Stoney Squaw Mountain, Banff N.P. (Brunton 1978);
southwest side Mount Norquay, Banff N.P. (H. F.
Coneybeare 1978); north side Banff Airfield, Banff
N.P. (Brunton 1978); 2.5 km W of Exshaw (Brunton
1978).
Western Cliff-brake is the smallest Canadian
Pellaea, rarely exceeding 10 cmin height. The Alberta
stations are on exposed hot limestone cliffs and rock
slopes with a southern or southwestern exposure. In
every station examined, Cheilanthes feei was a close
associate. It is typically found on the most exposed
cliffs and rock areas. It is locally very abundant in the
Bow Valley. The well-known Tunnel Mountain
station in Banff National Park supports hundreds of
thousands, if not millions, of plants. It is the largest
site known to date.
This species grows in very dense clumps in a “pin-
cushion” shape out of pocks and/or fissures in the
rock face. The fronds rarely extend out beyond the
surface of the rock face (as a water conservation
measure?). Despite the fact that it grows close to the
British Columbia border in Banff National Park and
1979 BRUNTON: PELLAEA IN ALBERTA 293
the Crowsnest Pass, there are no records for this
species W of the Continental Divide in Canada. The
increase in precipitation on the west side of the divide
is dramatic and appears to create unsuitable con-
ditions for P. occidentalis, even on sites which seem to
be otherwise quite suitable.
’ Banff
Mt ie ie
Cory Norquay\ _ KS
Jasper
National
Park
® Unpublished and/or new stations
A Previously published records
(Rigby and Britton. 1970)
kilometres
Kananaskis
Provincial
‘Three
A Sisters
Ona ae ae “ageonogtion
(pn a : ont : é
& * Sr
Outside of the Bow Valley, the Western Cliff-brake
is rare in Alberta. Being very inconspicuous, however,
it may well have been overlooked. It should be
watched for elsewhere in dry valleys along the eastern
slopes of the Rocky Mountains and on cliff areas in
the prairie (e.g., the Milk River canyon).
4
kilometres
6 8 10
\2,
CALGARY &
Rive’
i eae Suen
uF Feats
Whe
SL
FIGURE 5. Distribution of Pellaea occidentalis in Alberta.
294
THE CANADIAN FIELD-NATURALIST
Vol. 93
FIGURE 6. Typically dense clump of Pellaea occidentalis growing in hair-line fissure in open, south-facing limestone rock face.
South (5 km) of Canmore Alberta, | November 1977.
TABLE 1—Comparison of characteristics of Pellaea atropurpurea, P. occidentalis, and P. suksdorfiana. Source: 1, Rigby and
Britton (1970); 2, Tryon (1957); 3, This study
Characteristics
Chromosome no.
No. of spores per
sporangium
Frond length
Frond displacement*
Rachis color*
Stipes*
Pinnae*
Pinnae angle
to rachis
Site preference*
P. occidentalis
2n = 58
64 (small)
Maximum |[5 cm
Dense, thickly
clumped “pin
cushions” flush with
rock face
Golden to dark brown
Thin and brittle
old stipes
inconspicuous
Mitten-shaped and
sessile; margins
strongly reflexed
Perpendicular
Dry exposed
SW-facing limestone
P. suksdorfiana
2n = 116
32 (large)
Maximum 20 cm
Open and spreading,
extending out beyond
rock face
Reddish-brown to
brown-purple
Thicker and sturdy,
rarely sparsely
pubescent,
old stipes conspicuous
and often silvery color
Oblong-lanceolate and
(at least lower)
petioled; margins
strongly reflexed
Acute
Shaded cool, E-or
N-facing calcareous
cliffs by water
P. atropurpurea
2n = 87
32 (large)
Maximum 35 cm
Erect and spreading,
extending up and outward
from ledges far beyond
rock face
Purple to blackish-purple
Thick and sturdy,
heavily pubescent
old stipes abundant,
silvery color
Lanceolate to oblong-
lanceolate and
petioled; margins
slightly reflexed
Broad
Dry partially-shaded,
SW-facing limestone
Source
1
1,2
13
E253
*Feature of particular value for field identification.
1979
Discussion
There are many more stations for Pellaeain Alberta
than was previously thought. It isa common group on
the limestone cliffs and rock faces of the Bow Valley in
western Alberta. It is reasonable to expect that more
stations will be found in other drier mountain valleys
along the eastern slopes of the Rocky Mountains.
Pellaea suksdorfiana is the most frequently ob-
served Cliff-brake in Alberta, occurring uncommonly
in cooler shaded, more moist cliffs. Pellaea atro-
Purpurea appears to be locally common on sites
similar to that for P. occidentalis, both preferring hot
dry open limestone cliffs and rock faces. Pellaea atro-
purpurea prefers a more shaded and more heavily
fractured rock than does P. occidentalis.
With three of the four Canadian species occurring
in Alberta, the Pellaea flora of the province is
particularly rich. With so many stations growing in
close association in the Bow Valley, there is an
excellent opportunity to study the ecology of these
ferns and to search for (as yet unknown) hybrids. As
Tryon (1957) suggests that P. suksdorfiana may have
arisen as a fertile hybrid between P. atropurpurea and
P. occidentalis, this becomes an interesting possi-
bility.
Acknowledgments
My thanks to A. F. Tryon, M. Dumais, H. F.
Coneybeare, W.J. Cody, D.M. Britton, and D.
Jaques for information concerning their collections
and observations. The curators of CAN, DAO,
ALTA, and UAC provided access to these collections.
J. M. Gillett provided valuable assistance at CAN, as
did W. J. Cody and C. Frankton at DAO. In addition,
BRUNTON: PELLAEA IN ALBERTA
295
Cody loaned a number of specimens to me for
examination. D. M. Britton, University of Guelph,
reviewed an earlier manuscript and provided valuable
criticism. B. Wackerle typed several drafts of the
manuscript. I was accompanied in the field on several
occasions by D. Paton, H. F. Coneybeare, J. D.
Lafontaine, and B. Mitton. The late A. E. Porsild
provided important collection data early in the work
and encouraged this study. A particular vote of thanks
is owed my daughter Joni who hovered precariously
with me on numerous cliffs without complaint.
Literature Cited
Brunton, D. F. and J. D. Lafontaine. 1974. The distribu-
tion of Pellaea in Quebec and eastern Ontario. Naturaliste
Canadien 101: 937-939.
Hitchcock, C. L., A. L. Cronquist, and M. Ownbey. 1969.
Vascular plants of the Pacific Northwest. Volume 1.
University of Washington Press, Seattle.
Holmgren, P.K. and W. Keuken. 1974. Index herbar-
iorum, Part I. The herbaria of the world, 6th edition.
Regnum Vegetabile 92: 1-397.
Lafontaine, J.D. and D.F. Brunton. 1972. The Purple
Cliff-brake, Pellaea atropurpurea (L.) Link, in western
Quebec. Canadian Field-Naturalist 86: 297-298.
Rigby, S.J. and D. M. Britton. 1970. The distribution of
Pellaea in Canada. Canadian Field-Naturalist 84: 137-144.
Taylor, T. M. C. 1970. Pacific Northwest ferns and their
allies. University of Toronto Press, Toronto.
Tryon, A. F. 1957. A revision of the fern genus Pellaea
section Pellaea. Annals of the Missouri Botanical Garden
44: 125-193.
White, D. 1977. Rare plant survey: revisions. Trail and
Landscape 11: 22-25.
Received 29 November 1978
Accepted 31 March 1978
Characteristics of Peregrine Falcons Migrating through
Central Alberta, 1969-1978
DICK DEKKER
3819 — 112A Street. Edmonton, Alberta T6J 1K4
Dekker. Dick. 1979. Characteristics of Peregrine Falcons migrating through central Alberta. 1969-1978. Canadian Field-
Naturalist 93(3): 296-302.
In central Alberta from 1969 to 1978, 541 sightings of migrating Peregrine Falcons (Falco peregrinus) were recorded in spring,
and 34 in fall. Exclusion of suspected duplications gave 226 spring migrants and 27 fall migrants. In spring, adults arrived an
average of 9 d before the immatures. The great majority (86%) of spring migrants passed through the study area between 4
May and 23 May. Presence of adults peaked from 4 May to 7 May, of immatures from 12 May to 15 May. Fall sightings
occurred from 10 September to 4 October. In spring, males comprised 33% of the adult sample and 35% of the immatures
identified. Aggression between the sexes is suggested as a reason for the observed imbalance in the sex ratio. The proportion of
adults to immatures in spring was | to |. Spring migrants that were seen to leave the study area ina northerly direction did so
by soaring and high-altitude sailing. Strong head winds did not appear to deter some adults from migrating. Foraging activity
was seen to take place throughout the day and as late as | h after sundown.
Key Words: Peregrine Falcon, Falco peregrinus, migrations, Alberta.
Fall migration of northern Peregrine Falcons has
been studied along the coastal and Great Lakes
flyways (e.g., Hunt et al. 1975: Mueller and Berger
1961: Ward and Berry 1972), but very little infor-
mation has been recorded about their spring migra-
tion. Isolated spring sightings of peregrines have been
reported from across the continent, as evidenced by
scattered notes in American Birds. During a spring
study of migrating raptors along Lake Ontario only
four peregrines were seen in 108 d (Haugh and Cade
1966).
The paucity of data has led to speculation about the
routes, timing, and age structure of the peregrines
moving north. The fact that few subadults have been
observed on the breeding grounds suggested that
mortality of first-year birds is high. Cade (1960)
postulated that some immature peregrines may not
return to the Arctic and Subarctic until their second
year.
This paper provides information on the timing of
spring and fall migrations of peregrines through
Alberta, and the age, sex, and behavior of these
migrants.
Study Area and Methods
The study area is a crescent-shaped strip of land
situated on the rolling plains of central Alberta and is
roughly 15 X 2 km. Bordered by a marshy lake, it is
dominated by grain fields and pastures which hold
scattered pockets of Trembling Aspen (Populus
tremuloides) and willow (Salix spp.). Shallow depres-
sions collect melt-water in spring, attracting ducks
and shorebirds in large numbers during that season.
The study area was regularly visited from late
winter to late autumn to observe the hunting habits
of raptorial birds. During the 47-d period 15 April to
31 May, visits to the area averaged 27.9 d in each of
the 10 yr. During the 45-d period from 1 September to
15 October visits averaged 14.4 d. Days afield varied,
some lasting from sunrise until nightfall and others
spanning only 3 or 4h during the afternoon or
morning. Actual hours afield in spring were tabulated
only in the last 5 yr (Table 1).
In searching for peregrines, no set procedures were
followed. Vehicular travel in the study area was
limited, especially during wet periods. Most points
were accessibie only on foot, and 5-25 km were
covered by hiking each day.
Resting peregrines were detected by scanning the
landscape through 10-power binoculars. In calm
weather, the falcons sat on posts or stones and were
readily found, but strong winds induced them to
shelter on the ground. Very rarely were trees used as
perches. As a rule, to avoid flushing them, resting
peregrines were not approached closely. No attempts
at trapping or photography were made. Most falcons
were watched until they left of their own accord.
The majority of sightings were of flying peregrines.
To maximize chances of seeing hunting falcons, I
stayed near points where foraging peregrines had
previously been observed. Alarm reactions of prey
species, especially shorebirds, usually signalled the
appearance of a falcon. Often, while I was quietly
sitting in the open or in a parked car, the falcons
seemed to ignore my presence and approached boldly,
providing numerous close-range views. As the various
age and sex groups differ in appearance, and some
296
1979
birds show individual differences in plumage and
behavior, individuals sometimes could be recognized
with certainty: this helped me to minimize duplicate
counts.
The following procedure was used to reduce
possible duplications. When a bird identifiable as to
age and sex, for instance an immature female, was
observed on several consecutive days, each observa-
tion was recorded for the maximum count, but only
one sighting of an immature female was included in
the minimum count, unless there was evidence of
more than one immature female falcon present in the
study area. Sightings of unidentified peregrines were
included in the maximum count even though sex and
age were unknown, but not inthe minimum count if a
classified peregrine was known to be present in the
study area at the time.
Two or more interacting falcons were seen fre-
quently in aggressive encounters or in what appeared
to be cooperative hunting. Whenever this happened. I
could assess their numbers more accurately.
The passage of peregrines, like that of other
migratory birds, is seldom an evenly-spaced phe-
nomenon. There were periods when none were seen in
the study area, followed by days or parts of days when
several appeared in rapid succession. The rate of
arrival and departure influenced my ability to
calculate the number of individual peregrines seen
passing through the study area. I believe that the true
number is considerably less than the maximum count
presented but somewhat higher than the minimum
figure.
Results and Discussion
Numbers Sighted
Peregrines were sighted from 16 April to 30 May
(Figure |) and again from 10 September to 4 October
(Figure 2). From 1969 to 1978,541 peregrine sightings
were recorded in spring (Table 1), and 34 in fall
(Table 2). These include duplicate counts except the
most obvious ones. The minimum counts pooled for
10 yr are 226 in spring (Table 1) and 27 in autumn
(Table 2). From 1976 to 1978 both maximum and
minimum spring counts were much higher than
average, which can be attributed only partly to an
increase in field time (Table 1). These years were
characterized by early springs and low meltwater
levels, with few ponds available to ducks and waders,
which instead concentrated on the marshy lakeshore
bordering the study area. In consequence the falcons
hunted over a relatively narrow strip of land and were
more readily observed than in wet springs when they
foraged over much of the surrounding countryside.
Timing of Migrations
The earliest dates on which peregrines were
sighted in the study area in different years varied from
DEKKER: MIGRATING PEREGRINES IN ALBERTA
297
16 April to 9 May. These early arriving falcons were
always adults. The mean date of first arrivals was 27
April. First dates for the brown-backed immatures
varied from 27 April to 10 May. The mean first arrival
date for immatures was 5 May.9 d behind the adults.
The last spring dates on which adults were seen in
the study area ranged from 11 May to 27 May witha
mean last date of 19 May. Comparable dates for
immatures varied from 2! May to 30 May witha mean
of 25 May, 6 d after the adults. The great majority of
peregrines (86%) passed through the area from 4 May
to 23 May. Daily sightings showed a peak on 13 and 14
May. The peak of adult sightings occurred between 4
and 7 May, 8 d before the peak of immatures which
fell between 12 and 15 May (Figure 1).
The limited data from the autumn show sightings
from 10 September to 4 October with a peak on 23
September (Figure 2).
Subspecies and Plumage Variation
White (1968) recognized three subspecies of Pere-
grine Falcons in North America. Falco p. anatum
formerly nested in nearly all suitable habitat south of
the tundra except the northwest Pacific coast where F.
p. pealei occurs. Falco p. tundrius inhabits the Arctic
and Subarctic. During this study no representatives of
F. p. pealei were identified. Most of the migrants seen
could not be assigned to any subspecies. A few falcons
that appeared typical, in some respects, of F. p.
anatum were seen during the early part of each spring
season, while typical examples of F. p. tundrius were
among the last peregrines to pass through. The _
criterion used for field identification of the two
subspecies was dorsal coioration which could best be
appraised when the bird was in flight. The blue-gray
color of adult F. p. anatum was lightest on the rump
and darkened to nearly black on the head, tail, and
wingtips. This variegated color pattern of F. p.
anatum was also noted by Beebe (1974). When in
flight, typical F. p. tundrius adults appeared evenly |
blue-gray on the entire dorsal surface. They usually
looked smaller and slenderer than typical F. p.
anatum birds of the same sex.
All adult falcons that could be observed through a
20-power scope at reasonably close range showed a
very light, unmarked upper breast. Occasionally there
was a pinkish wash on the remainder of the ventral
surface. Dark barring and spotting on the belly and
flanks varied from bold to fine, and from dense to
widely spaced. Greater diversity of pigmentation and
wider extremes in light and dark types were found
among the first-year birds. Few spring immatures
were streaked on the entire undersurface as they are in
the preceding autumn. The majority showed a
varying amount of unmarked yellowish-white on the
breast, which made them resemble adults. To describe
these falcons properly, a good view of the dorsal color
Vol. 93
THE CANADIAN FIELD-NATURALIST
298
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1979 DEKKER: MIGRATING PEREGRINES IN ALBERTA 299
on 8 8 6 G 8
ul
FIGURF 1. Maximum numbers of peregrines sighted in spring. Data pooled for 10 years. 1969-1978.
10
PEREGRINES
5 10 15 20 Zs) 30 1 5 10 15
SEPTEMBER OCTOBE
FIGURE 2. Maximum numbers of peregrines sighted in autumn. Data pooled for 10 years. 1969-1978.
300
had to be obtained. Their brown backs ranged in tone
from blackish-brown to sandy. The pale brown F. p.
tundrius immatures were easily mistaken for Prairie
Falcons (Falco mexicanus), which were only rarely
seen in the study area during spring and fall. In fall,
second-year peregrines had attained adult coloration
although their wings and backs retained some
brownish feathers.
Of interest was the occurrence in the study area of
“red-tailed” peregrines. About one-third of the adults
showed a buffy or brownish cast on the upper tail.
Also some immature falcons had “red” tails. The color
varied from dark reddish brown to a strikingly pale
buffy-orange. To my knowledge, there are no records
in the literature of such “red-tailed” peregrines.
Age Ratios
During 10 springs, 102 immature peregrines were
recorded among a total of 202 falcons classified as to
age. This age ratio cannot be considered representa-
tive of the population as a whole. Shor (1970, p. 56)
reported that peregrine populations contain more
adults than first-year individuals “even at the time of
the fall migration.” This difference should be even
more pronounced in spring, as Enderson (1969)
calculated a 70% mortality among first-year birds
from band returns.
The very small sample of fall data obtained
included 14 immatures and 6 adults. Fall counts from
coastal migration routes indicate immature to adult
ratios of 4 to | and 6 to! (Berry 1971; Ward and Berry
1972: Hunt et al. 1975). These ratios suggest dif-
ferential migratory behavior among the age classes.
Berry (1971) argued that adults, being more experi-
enced in foraging, could fly directly to their wintering
grounds.
My observations suggest that during spring both
age classes hunt the same prey and ina similar way.
Hunting success ratios of adults and of immature
falcons migrating through central Alberta during
spring are not significantly different (Dekker, un-
published data). Hence, the observed prominence of
first-year young in central Alberta in spring cannot be
explained as a consequence of different hunting
ability as suggested by Berry (1971) for fall migrants.
The majority of peregrines known to linger in the
study area, however, were in immature plumage. The
duplication rate of immatures was 2.3 (102 to 239),
that of adults was 1.6 (100 to 167). It appeared that
adult peregrines on their way north through central
Alberta were less inclined to tarry on attractive
feeding grounds than the immature birds.
Sex Ratios
The smallest male peregrines are about two-thirds
the size of the largest females (Godfrey 1966). Hunt et
al. (1975), however, have recognized the difficulty of
THE CANADIAN FIELD-NATURALIST
Vol. 93
accurately estimating sex ratios on the basis of
sightings. I agree that the size of a peregrine cannot be
judged reliably while the bird is resting unless it can be
compared to another bird. In flight, however, a male
peregrine usually appears slenderer than a female.
Small to average-sized males and large to average-
sized females are readily distinguished when seen in
flight by an experienced observer. Additionally,
foraging habits may provide clues and assist in
classification. For example, males are more given
than females to pursue and swoop repeatedly at
fleeing sandpipers. About 85% of the peregrines
included in the minimum count and classified to sex
were observed under conditions that allowed me to
make such classification with confidence. An element
of doubt is present in about 15% of cases when
individual variation in size made the sexes look alike.
Rather than assigning these puzzling birds to the
unidentified category, I classified them on the basis of
impressions. As only one observer was involved these
impressions may be expected to show a degree of
consistency. Sex ratios obtained for adult (33%) and
immature peregrines (35%) in fact show a striking
similarity.
Adult males are reported to be most elusive along
the fall migration routes. Only eight out of 639
peregrines trapped on Assateague Island from 1939 to
1971 were identified as adult males (Ward and Berry
1972). No adult males were among 250 peregrines
captured on the Texas beaches from 1952 to 1973
(Hunt et al. 1975), and only one among 20 classified in
fall in this study. Percentages of immature male
peregrines trapped along the major fall migration
routes along the Texas and Maryland coasts respec-
tively were reported as 23.4% (Hunt et al. 1975) and
30% (Ward and Berry 1972).
Hunt et al. (1975) claim that there is no obvious
reason why immature sex ratios should be un-
balanced. The observed preponderance of females in
the Texas survey may reflect differential trap-
response and/or foraging habits. The males are
believed to be able to exploit inland populations of
small birds better than are the larger females, and are
less often seen on the beaches. In my opinion,
aggression between sexes may force the smaller males
away from localities where female peregrines congre-
gate. I frequently saw females chase males. Males were
observed to carry their prey long distances before
alighting to eat it. Two were observed to soar to great
altitudes and feed while on the wing. By contrast,
females routinely consumed their prey at the site of
capture. On two occasions an adult female was seen to
rob an immature male of his freshly caught prey.
Attempts at stealing each other’s prey were seen also
between immature females and between immature
and adult females.
1979
Weather and Daily Routine
The influence of weather on the movements of
migrating raptors in spring and fall has been discussed
in detail by Mueller and Berger(1961) and Haugh and
Cade (1966). They concluded that the majority of
raptorial birds migrate when tail winds are blowing.
Ward and Berry (1972) and Hunt et al. (1975)
correlated highest numbers of peregrines in fall with
days of heavy cloud, light tail winds, and low
temperatures. Those conditions usually accompanied
the influx of some falcons into my study area in fallas
well as in spring. The birds were easily located then as
they rested on exposed perches (fence posts) or hunted
at low elevations. Mueller and Berger (1961) have
remarked that hawks have rarely been observed in the
process of departing on migration. During this study
many resting falcons were watched until they flew
away of their own accord. In spring they consistently
left the study area in a north-northwest direction.
Such departures were seldom noticed on days of
heavy cloud and low temperatures.
Of particular interest was the observation that light
to moderate head winds at ground level did not
prevent some falcons from departing. They did not
migrate close to the ground, as stated by Fischer
(1967), but maintained high altitudes attained by
soaring. Brown and Amadon (1968, p. 65) write that
“falcons usually fly rather than sail or soar” while
migrating. Both migrating and foraging falcons
observed in this study not only soared during
optimum conditions of sun and wind, but also on
windy days when skies were cloudy, and even when
light rain was falling. When nearly straight overhead
of an observer, some of these soaring falcons were just
visible in 10-power binoculars. On two occasions
soaring peregrines were seen to enter the cumulus
clouds that top thermal currents.
Falcons that were seen to leave the study area ona
steady northerly course on spring days when head
winds were blowing, habitually winged upwind for a
short distance before beginning to soar. During their
circling ascent the falcons drifted downwind until they
flexed their wings and glided through the wind
rapidly. Even head winds gusting to50 km per hour at
ground level did not deter them from high-altitude
travel. All falcons observed to migrate during these
contrary airflows were adults. First-year falcons that
were seen to travel northward did so when tail winds
were blowing or when head winds were only light.
Most observed departures of adults and immatures
took place during mid-morning after the air had
warmed sufficiently to allow the formation of rising
air currents.
During the remainder of the day some migrating
peregrines were located when they descended to
attack prey species such as feeding shorebirds. After
DEKKER: MIGRATING PEREGRINES IN ALBERTA 301
consuming a kill, a peregrine would resume flight by
soaring to great heights and sailing away on slightly
flexed wings. Such interruption and resumption of
daytime migratory travel was observed from 11:00 to
18:00 hours. In spring, during the late afternoon and
early evening, falcons sometimes flew in from a
southerly direction as if they were just arriving. Aftera
rest, these falcons began hunting in the hour preceding
sundown. Several immature falcons were seen chasing
and capturing shorebirds shortly before dusk. One
immature female falcon that had been watched from
late afternoon to nightfall began attacking ducks that
flew inland from the lake one hour after sundown.
Through binoculars, the birds were just visible,
silhouetted against a cloudless sky. Crepuscular
hunting was also observed by Beebe (1960), Clunie
(1976), Fischer (1967), and Rudebeck (1950-1951).
Late foraging activity may explain the relative scarcity
of falcons on the Texas beach at sundown, which
prompted Enderson (1965) to speculate on the likeli-
hood of nocturnal migrations.
During this study, several falcons were observed
roosting at nightfall. They were sitting on posts
surrounded by water or on shoreline rocks. Of three
adult peregrines that were seen roosting at night only
one was still on its post when it was checked at sunrise.
It remained there until 10:00 hours when it flew
upwind, began to soar and sailed northward.
I have seldom found evidence of hunting at dawn.
An hour after sunrise was the usual start of foraging
flights. When a large food item such as a duck was
captured, the falcon would eat for about 45 min and
rest until mid-morning. A second meal of the prey
remains was frequently taken before the falcon flew
away, soared to great height, and glided northwards
until lost to view. Some spring migrants that soared
and sailed northwards during mid-morning were seen
to stoop at prey species that flew below them: it
appeared that these peregrines did their foraging
along the way.
NOTE: Formerly the study area was part of the
summer range of peregrines that nested along rivers to
the south and west about 100 kmaway. By 1969, at the
start of this survey, that central Albertan population
of F. p. anatum was all but extirpated (Dekker 1967).
In 1977 and 1978 one or two peregrines frequented the
area during the summer months. In 1978 a first-year
male, recognizable by his banded feet, damaged tail,
and lack of shyness, was often seen from April to late
August. These birds were believed to have originated
from the captive breeding program sponsored by the
Canadian Wildlife Service and the Provincial Fish
and Wildlife Division. All known and_ suspected
sightings of these birds have been eliminated from the
tabulations in this paper.
302
Acknowledgments
During the springs of 1974 to 1978, this study was
financially supported by the Alberta Department of
Recreation, Parks and Wildlife under the direction of
Wiiliam Wishart, Head of Wildlife Research. I thank
D. A. Boag for his valuable suggestions for revising
the manuscript. I am also grateful for the constructive
criticism offered by A. J. Erskine, D. V. Weseloh, and
adie Cade:
Literature Cited
Beebe, F. L. 1960. The marine peregrines of the Northwest
Pacific Coast. Condor 62: 145-189.
Beebe, F. L. 1974. Field studies of the Falconiformes of
British Columbia. British Columbia Provincial Museum.
Occasional Paper Series Number 77. 163 pp.
Berry, R.B. 1971. Peregrine falcon population survey.
Assateague Island. Maryland. fall 1969. Raptor Research
News 5: 31-43.
Brown, L. and D. Amadon. 1968. Eagles. hawks and
falcons of the world. Hamlyn House. 945 pp.
Cade, T. J. 1960. Ecology of the peregrine and gyrfalcon
populations in Alaska. University of California Publica-
tions in Zoology 63: 151-290.
Clunie, F. 1976. A Fiji peregrine in an urban environ-
ment. Notornis 23: 8-28.
Dekker, D. 1967. Disappearance of the peregrine falcon as
a breeding bird of a river valley in central Alberta.
Blue Jay 25: 175-176.
Enderson, J. H. 1965. A breeding and migration survey of
the peregrine falcon. Wilson Bulletin 77: 327-339.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Enderson, J.H. 1969. Peregrine and prairie falcon life
tables based on band recovery data. /nm Peregrine
populations — their biology and decline. Edited by
J.J. Hickey. pp. 505-509.
Fischer, W. 1967. Der Wanderfalk. Die Neue Brehm
Bucherei. Ziemsen Verlag. 152 pp.
Godfrey, W.E. 1966. The birds of Canada. National
Museum of Canada. Bulletin 203. 428 pp.
Haugh, J.R. and T.J. Cade. 1966. The spring hawk
migration around the southeastern shore of Lake Ontario.
Wilson Bulletin 78: 88-110.
Hunt, W. G., R. R. Rogers, and D. J. Slowe. 1975. Migra-
tory and foraging behavior of peregrine falcons on the
Texas coast. Canadian Field-Naturalist 89: 111-123.
Mueller, H. C. and D. D. Berger. 1961. Weather and fall
migration of hawks at Cedar Grove. Wisconsin. Wilson
Bulletin 73: 171-192.
Rudebeck, G. 1950-1951. The choice of prey and modes of
hunting of predatory birds with special reference to their
selective effect. Oikos 2: 65-88: 3: 201-231.
Shor, W. 1970. Peregrine falcon population dynamics
deduced from band recovery data. Raptor Research News
4: 4959.
Ward, F. P. and R. B. Berry. 1972. Autumn migrations of
peregrine falcons on Assateague Island. 1970-1971.
Journal of Wildlife Management 36: 484-492.
White, C. M. 1968. Diagnosis and relationships of the
North American tundra inhabiting peregrine falcons.
Auk 85: 179-191.
Received 30 October 1978
Accepted 19 March 1979
Notes
Extralimital Record of a Narwhal (Monodon monoceros)
in Hall’s Bay, Newfoundland
BORA MERDSoy,! JON LIEN, and ANNE STOREY
Department of Psychology, Memorial University of Newfoundland, St. John’s, Newfoundland AIC 5S7
\Present Address: 7 Appledore Place, St. John’s, Newfoundland A1lB 2W8
Merdsoy, Bora, Jon Lien, and Anne Storey. 1979. Extralimital record of a Narwhal (Monodon monoceros) in Hall’s
Bay, Newfoundland. Canadian Field-Naturalist 93(3):
Key Words: Narwhal, Monodon monoceros, Newfoundland,
On 1 March 1978 a Narwhal (Monodon
monoceros) was seen from a small boat in Hall’s Bay,
Newfoundland (49°30’N, 56°00’W). Pack ice, several
miles wide, had jammed across the head of the fjord
and extended into the bay so that the animal, along
with four Humpback Whales (Megaptera novae-
angliae), was confined to an area 8 X4km.
Humpback Whales were observed in the area earlier
in the winter, but the Narwhal was not seen until
February. Both local and arctic ice conditions were
unstable within the bay, but the entrance to the bay
remained blocked so that the Humpback Whales and
the Narwhal were unable to leave Hall’s Bay until 6
April. At this time the pack ice, loosened several days
earlier by an ice-breaker, drifted to open water
enabling the whales to escape. No sightings of the
Narwhal or Humpback Whales were made after they
left Hall’s Bay although repeated air surveillance was
attempted (Cyril Pelly, Springdale, Newfoundland,
personal communication).
Hall’s Bay contains | m of fresh water at the surface
which results in rapid formation of local ice over the
entire bay. During the period the whales were
confined to the bay the area of open water decreased
irregularly. For a period of 3 wk the only open water
was 1.6 km NW of Mansfield Head, which varied in
size from 120 X 45 m. This confinement allowed for
close observation and photography of the Narwhal
(Figure 1).
The Narwhal was a dark gray color, 2-3 m long
with a tusk of approximately 15-20 cm, indicating
that it was a male. Generally the Narwhal surfaced in
the open water during periods the Humpback Whales
were below the surface. In addition to such time-
sharing behavior, spatial separation was maintained,
as the Humpback Whales typically would follow the
edges of the open water while the Narwhal restricted
its surface activities to the center. Although ice forced
the Narwhal into proximity with human observers, it
remained timid throughout its confinement typically
maximizing its distance from observers. Food species
is)
i)
303-304.
sighting, range, geographical distribution, pack ice.
including Atlantic Herring (Clupea harengus),
Rainbow Smelt (Osmerus mordax), and Capelin
(Mallotus villosus) were plentiful in the area and
hydrophone recordings indicated hunting activity by
the animal (Ford and Fisher 1978). Nosurface sounds
were heard from the Narwhal.
The range of the Narwhal in the Canadian Eastern
Arctic extends south to about 66°N (Banfield 1974).
Our sighting was approximately 1800 km south of this
at 49°30’N. Our report is the second sighting of a
Narwhal in Newfoundland in the last decade. The first
extralimital record of a mature male Narwhal, found
washed on a beach in White Bay in 1969, was reported
by Mercer (1973). Several sightings of white whales
reported in Newfoundland waters (Sergeant and
Fisher 1957; L. M. Tuck, St. John’s Newfoundland,
personal communication), could be either Belugas
(Delphinapterus leucas) or Narwhals. The south-
eastward movement of arctic ice in the late winter
brings Harp (Phoca groenlandicas) and Hooded
(Cystophora cristata) Seals in large numbers to
Newfoundland’s coast. Occasionally Bearded Seals
(Erignathus barbatus), Ringed Seals ( Phoca hispida), |
Walrus (Cdobenus rosmarus), and Polar Bears ( Ursus
maritimus) have drifted south with the ice (L. M.
Tuck, personal communication).
Narwhals are reported to winter in the Baffin Bay
area (Mansfield et al. 1975) and it is probable that, on
occasion, they follow ice southward. This could
account for the present record, but may not account
for the June 1969 record (Mercer 1973) unless that
animal came with the ice and survived in
Newfoundland through the spring. Because it is
possible to mistake the finless dorsum of an immature
Narwhal for that of a seal, it may be that more
Narwhals frequent the sparsely inhabited shores of
Newfoundland than are being recognized.
We gratefully acknowledge the help of Francis Hull
and Cyril Pelly of Springdale, Newfoundland, during
the period of our work in Hall’s Bay.
304 THE CANADIAN FIELD-NATURALIST
FIGURE |. The Hall’s Bay Narwhal during the last week of its confinement. The tusk length was approximately 45 cm, which
aged the animal at over 2 yr. Photographed by Duncan Findlayson.
Literature Cited
Banfield, A.W. F. 1974. The mammals of Canada.
University of Toronto Press, Toronto. 438 pp.
Ford, J. K. B. and H. D. Fisher. 1978. Underwater acou-
stic signals of the Narwhal (Monodon monoceros).
Canadian Journal of Zoology 56: 552-560.
Mansfield, A. W., T. G. Smith, and B. Beck. 1975. The
Narwhal, Monodon monoceros, in eastern Canadian
waters. Journal of the Fisheries Research Board of
Canada 32: 1041-1046.
Mercer, M.C. 1973. Observations on distribution and
intraspecific variation in pigmentation patterns of
odontocete Cetacea in the western North Atlantic.
Journal of the Fisheries Research Board of Canada
30: 1111-1130.
Sergeant, D.E. and H.D. Fisher. 1957. The smaller
Cetacea of eastern Canadian waters. Journal of the
Fisheries Research Board of Canada 14: 83-115.
Received 19 September 1978
Accepted 30 January 1979
First Canadian Record of the Black Buffalo
(Osteichthyes: Catostomidae)
E. J. CROSSMAN! and S. J. NEPSzyY2
'Department of Ichthyology and Herpetology, Royal Ontario Museum, Toronto, Ontario M5S 2C6
2Ontario Ministry of Natural Resources, R.R. #2, Wheatley, Ontario NOP 2P0
Crossman, E. J. and S.J. Nepszy. 1979. First Canadian record of the Black Buffalo (Osteichthyes: Catostomidae).
Canadian Field-Naturalist 93(3): 304-305.
Ictiobus niger was captured in the Canadian waters of Lake Erie on 20 June 1978.
Key Words: Black Buffalo, Jctiobus niger, first Canadian record, Lake Erie, geographical distribution.
A specimen of J/ctiobus niger, the Black Buffalo,
one of the larger deep-bodied suckers, was captured in
the Canadian waters of Lake Erie on 20 June 1978. It
had been taken in a commercial trap net set by F.W.
1979
NOTES
305
TABLE 1—Comparison of characters in the three species of Ictiobus
Character Ictiobus cyprinellus
Ictiobus bubalus Ictiobus niger
Body Robust, slightly com-
pressed, back not highly
elevated
Usually 24-28
Large, oblique
Dorsal rays
Mouth
Lips Thinner than others, only
faintly striate, upper
about level with lower
margin of orbit
Distance mandibular
symph. to tip of maxilla
Upper jaw About as long as snout
Gill rakers post. face
first arch
Lower pharyngeal arch
Nearly 100
Thin, more than twice as
high as wide, teeth weak
More slender, but thicker
than /. b., less elevated
and less sharpened than J. b.
Deeper and narrower than
Ln., highly elevated, mid
dorsal ridge sharper than
others
Usually 27-30
Smaller, less oblique than Less oblique than /. c.,
I. c., more inferior than larger and less inferior than
Ln. ae:
Thicker, more or less Thicker than /.c., more
coarsely straite, upper or less coarsely striate,
far below lower margin upper far below lower
of orbit margin of orbit
Only slightly greater than As much as twice diam. of
diam. of orbit in adults orbit in adults
Distinctly shorter than Shorter than snout
snout
Fewer than 60
Usually 30
Fewer than 60
Heavy. about as wide as
high, teeth strong
Heavy. about as wide as
high, teeth strong
Krause and Sons Fisheries. Leamington, Ontario, in
Pigeon Bay. west of Point Pelee (41°56’50’N.
82°31'40”W). The specimen was turned over to
Nepszy at the Ontario Ministry of Natural Resources
Research Station Wheatley, Ontario, and forwarded
to Royal Ontario Museum, frozen, as an unidentified
sucker. possibly [ cyprinellus. After fixation in
formalin, and preservation in ethy] alcohol the buffalo
was 630 mm in total length. and weighed 3.56 kg.
Preliminary examination. including cranial fon-
tinelles. clarified that it was [ctiobus but that it was
not J. cyprinellus. the commonest species and the only
species previously reported as occurring in Canada.
Morphologically it seemed to resemble both 1
bubalus and I. niger and was somewhat suggestive ofa
hybrid. Neither of those species. however. is common
in Lake Erie and published characteristics used to
separate the two species are difficult to interpret.
Ictiobus niger is basically a species of the Missis-
sippi River system and occurs from the Ohio River
and its tributaries south to Mexico. It has. however.
been said to be native to southern Lake Michigan
(Hubbs and Lagler 1964) and therefore in the Great
Lakes system. Moore (1968) listed 1. niger but not 1.
bubalus from Lake Erie. and Hubbs and Lagler (1964)
specified earlier that there was one report only of /.
niger, and gave no Great Lakes record for J. bubalus.
Trautman (1957) gave no records for the Ohio waters
of Lake Erie for either species. Trautman did indicate.
however (under /. bubalus), that specimens had been
caught in Sandusky Bay. Lake Ene. which had been
identified as hybrids between [. cyprinellus and I.
bubalus or I. niger.
So few specimens of ZI/ctiobus other than 1
cyprinellus are seen from the Great Lakes that the
characteristic features of larger specimens of /.
bubalus and I. niger are poorly known. A comparison
of the features of the three species is given in Table 1.
The supposed hybrids may have represented the
presence of I. bubalus or I. niger in Lake Erie in the
past.
After detailed examination of the characteristics,
including the cranial bones, of this specimen, G. R.
Smith and R. M. Bailey of the Museum of Zoology.
University of Michigan confirmed that the
specimen represents Ictiobus niger. Therefore the
specimen is not only a recent corroboration of the
occurrence of the species in Lake Erie but also
represents a species new to the fauna of Canada. It has
been added to the collection of the Royal Ontario
Museum (ROM 34562).
We once more thank Rudy Krause for his vigilance
and for turning over specimens suggestive of some-
thing new. We thank Erling Holm. and G. R. Smith
and R. M. Bailey for their help in the identification of
the specimen.
Literature Cited
Hubbs, C. L. and K. F. Lagler. 1964. Fishes of the Great
Lakes region. University of Michigan Press. 213 pp.
Moore, G. A. 1968. Fishes. Jn Vertebrates of the United
States. McGraw-Hill Inc. pp. 21-165.
Trautman, M.B. 1957. The fishes of Ohio. Ohio State
University Press. 683 pp.
Received 13 December 1978
Accepted 30 January 1979
306
THE CANADIAN FIELD-NATURALIST
Vol. 93
Thaspium trifoliatum (Meadow-parsnip) in Canada
P. W. BALL
Department of Botany, Erindale College, University of Toronto, Mississauga, Ontario LSL 1C6
Ball, P. W. 1979. Thaspium trifoliatum (Meadow-parsnip) in Canada. Canadian Field-Naturalist 93(3): 306-307.
Thaspium trifoliatum (Meadow-parsnip) is reported from a locality near Amherstburg, Essex County, southern Ontario.
Although reported from Canada on a number of previous occasions, all the earlier records appear to be based on confusion
with the very similar Zizia aptera. The differences between T. trifoliatum and Z. aptera are discussed and it is noted that, in
addition to the fruit characters usually used to distinguish between them, the width of the petiole also appears to be diagnostic.
Key Words: Thaspium trifoliatum, Umbelliferae (Apiaceae), Essex County, Ontario.
Thaspium trifoliatum (Meadow-parsnip) has been
recorded from Canada on a number of occasions.
Indeed, Macoun (1883) states that the species is
abundant in the prairie provinces. Subsequent
investigations have shown that all of these records are
erroneous. For example, Scoggan (1957) makes it
quite clear that all Manitoba records of T. trifoliatum
should be referred to Zizia aptera. Soper (1949) lists
T. trifoliatum from southern Ontario, this record
apparently being based on a specimen from Caradoc,
Middlesex County, collected by J. Dearness in 1891,
and now in the herbarium of the Biosystematics
Research Institute, Ottawa (DAO). This specimen
was subsequently redetermined by W. J. Cody as Z.
aptera, and, although the specimen is immature, it
does appear to be this species. Boivin (1967) did not
include T. trifoliatum in the Canadian flora and, ina
personal communication, has stated that he has not
traced any reliable records since then.
The discovery of a location for 7. trifoliatum in
southern Ontario seems therefore to be the first
authentic record for this species in Canada. The
locality is a woodlot about 3 km E of Amherstberg in
Essex County. The dominant trees were Quercus alba
(White Oak), Ostrya virginiana (Hophornbeam),
Carya ovata (Shagbark Hickory), and Fraxinus
pennsylvanica (Green Ash). The shrub layer was fairly
dense with Zanthoxylum americanum (Prickly Ash),
Rhus aromatica (Fragrant Sumach), Viburnum
rafinesquianum, and Crategus spp. (hawthorn)
prominent in the vicinity of the Thaspium. The stand
also contained Geum vernum, another species that is
rare in Ontario and largely restricted to Essex County.
The population of 7. trifoliatum consisted of over 50
individuals so does not appear to be in immediate
danger of extinction; however, any major disturb-
ance of the woodlot could seriously jeopardize the
survival of this species in Canada.
Thaspium trifoliatum and Z. aptera are remar-
kably similar in appearance, so that confusion
between these two species is not surprising. When in
fruit they can be readily distinguished by two
characters. Zizia aptera has fruits with narrow ridges
(Figure 1A) and the central fruit of each subumbel is
sessile; T. trifoliatum has fruits which are distinctly
winged (Figure 1B) and which are all distinctly
pedicelled. Unfortunately these differences cannot be
easily observed when the plants are in flower. The
wings on the fruits of T. trifoliatum are not apparent
until a fairly late stage in fruit development. The
presence of a central sessile flower in each subumbel
can be readily observed once all the flowers are open,
but when the inner flowers of each subumbel are in
bud it is difficult to determine whether the central
flower is sessile or pedicelled. An additional character
which is useful, although not completely reliable, is
the width of the petiole of the cauline leaves (Figure
1C, D). In Z. aptera the petioles of the lower leaves on
the flowering stem are 8-16 mm wide and almost forma
sheath around the stem, whereas in 7. trifoliatum the
petioles of the lower leaves are 4-8 mm wide and
scarcely enclose the stem except at the node. The
petioles of the upper leaves of both species are
somewhat narrower, 5-10 mm wide in Z. aptera and
3-5 mm wide in T. @rifoliatum, but if leaves are
compared from similar regions of the stem then the
petioles of Z. aptera are always wider than those of 7.
trifoliatum.
Thaspium trifoliatum is variable in flower color.
Var. trifoliatum, which is the predominant variety in
the Appalachian region, and which also occurs rarely
elsewhere, has purple flowers. Var. flavum, which has
yellow flowers, is more western in distribution
occurring as far west as Minnesota, Arkansas, and
Kansas, but it also occurs rarely in the east. The
Ontario material is referable to var. flavum.
Specimen Examined
Essex County. 3 km (2 mi) E of Amherstberg. 10
June 1975. P. W. Ball 14475 (TRTE, DAO) (Flower,
young fruit). Anderdon Township 3 km (2 mi) NE of
Amherstberg on east side of Concession II. 26 May
1977. P.M. Catling (TRT) (Flower). June 1977.
P. M. Catling (TRT, TRTE, CAN). (Fruit).
ISD
NOTES
307
G
FiGure |. Fruit and upper leaves of Zizia aptera (A, C) and Thaspium trifoliatum (B, D).
Acknowledgments
My thanks to J. McNeill for checking the identity of
some of the specimens and to P. Catling for collecting
some specimens in fruit. This work was supported by
National Research Council of Canada Grant A6494.
Literature Cited
Boivin, B. 1967. Enumération des plants du Canada. III.
Naturaliste Canadien 93: 583-646.
Macoun, J. 1883. Catalogue of Canadian plants. Part 1.
Montreal. 192 pp.
Scoggan, H. J. 1957. Flora of Manitoba. National Muse-
um of Canada Bulletin 140. 619 pp.
Soper, J. H. 1949. The vascular plants of southern Ontario.
Toronto. 95 pp.
Received 25 May 1978.
Accepted 19 January 1979.
308 THE CANADIAN FIELD-NATURALIST
Vol. 93
Largest Gray Wolf Skulls Found in Alberta
JOHN R. GUNSON! and RONALD M. NOWAK2
\Alberta Fish and Wildlife Division, 6909-116 St., Edmonton, Alberta T6H 4P2
2Office of Endangered Species, U.S. Fish and Wildlife Service, Washington, D.C. 20240
Gunson, J.R. and R.M. Nowak. 1979. Largest Gray Wolf skulls found in Alberta. Canadian Field-Naturalist
93(3): 308-309.
Measurements of greatest length and zygomatic width of five large skulls of Canis lupus from Alberta are reported. Three, all
from the range of C. /. occidentalis, are larger than the previous largest specimen, also occidentalis.
Key Words: Gray Wolf, skull measurements, Canis lupus, size.
According to Goldman (1944) the largest sub-
species of the Gray Wolf (Canis /upus) are found in
Alaska and northwestern Canada. With respect to
measurements in millimetres of greatest length (g.1.)
and zygomatic width (z.w.) of skull, the four largest
individuals that he listed (pp. 490-497), all males, were
the following: United States National Museum 9001,
C. | occidentalis, Fort Simpson, Mackenzie, g.l.
292.8, z.w. 156.5; Royal Ontario Museum 33-9-20-5,
C. |. pambasileus, White River, Yukon, g.1. 293.7, z.w.
151.5: Royal Ontario Museum 31-2-16-2, C. 1.
pambasileus, White River, Yukon, g.]. 292.8, z.w.
149.0; University of California Museum of Vertebrate
Zoology 31043, C. /. columbianus, Iskut Summit,
British Columbia, g.1. 288.9, z.w. 155.0. On the basis
of both measurements, the specimen from Fort
Simpson, which was collected in 1869, has been until
now the largest reported skull of a North American
Gray Wolf.
Several specimens of male C. /upus recently
collected in Alberta have skull dimensions that
approximate or surpass those of the above-mentioned
material (Table 1). The largest of all, number
Z78.104.1 in the collection of the Provincial Museum
of Alberta (formerly 66-18 of the Alberta Fish and
Wildlife Division), was taken at Roche Lake in the
Swan Hills area, about 155 km northwest of Edmon-
ton. Two others, Z78.104.2 and Z78.104.3, taken
during 1972-1974 in northwestern Alberta were also
larger than the Fort Simpson wolf. All three were
taken in areas within the original range of C. /.
occidentalis, as was Z78.104.4. Specimen Z78.104.5
was collected in southwestern Alberta from an area
within Goldman’s original range of C. /. irremotus.
Determination of the taxonomic position of that
specimen and others recently collected from even
more southerly locations in Alberta awaits the
collection of additional data.
Several extensive regional collections of North
American Gray Wolves have been assembled in recent
decades, but with the exception of the Alberta Fish and
Wildlife Division collection (12 measurements on
each of 468 skulls), none have been thoroughly
analyzed. One of us (Nowak) made cursory examina-
tions of a collection from western Canada in the
possession of the University of British Columbia and a
collection from Alaska in the possession of the
University of Alaska. No skulls were found that
surpassed in size the larger of the above-described
specimens, but one of the Alaska skulls of C. /.
pambasileus may be the largest ever recorded for that
state. The specimen (University of Alaska 40682) was
TABLE 1—Specifics of five large male Canis /upus from Alberta, Canada. PMA = Provincial Museum of Alberta:
AFW = Alberta Fish and Wildlife Division
Specmontie Greatest Zygomatic
: Date of length width
PMA AFW Location collection (mm) (mm)
Z78.104.1 66-18 54°46’N,114°55’W 30 Jan. 1966 304.5 154.8
Z78.104.2 V441036 55° 4’N,119°50’W 13 Feb. 1973 297.6 158.0
Z78.104.3 V441097 54°43’N,113°17’W Feb. 1974 288.8 162.5
Z78.104.4 V441057 54°48’N,119°20’W 27 Feb. 1973 292.6 155.8
Z78.104.5 V441016 $1°27’N,114°50’W 28 Feb. 1973 285.3 160.4
1979
collected in 1966 in the Wood River drainage of
central Alaska, and its measurements (taken to the
nearest millimetre) are g.l. 292, z.w. 153.
Mean greatest length and zygomatic width for 83
adult male C. /upus taken in Alberta during
1965-1966 to 1977-1978 were 275.6 (range 251.3 to
304.5) and 150.7 (range 128.6 to 162.5). The following
data on series of North American wolf skulls (Nowak
1973) are offered for purposes of comparison: 233
Recent male C. /upus from throughout northern and
western North America, mean g.l. 259.6 (range
235-293), mean z.w. 141.1 (range 126-164); 20 male C.
|. pambasileus, mean g.1. 271.4 (range 248-288), mean
z.w. 145.4 (range 130-154): 62 C. dirus(unsexed) from
late Pleistocene deposits at Rancho La Brea, Cali-
fornia, mean g.l. 294.8 (range 258-316), mean z.w.
163.3 (range 148-177). In comparisons of skull
dimensions of wolves from several areas in western
North America, Skeel and Carbyn (1977) recorded
largest mean g.|. and z.w. in specimens from Prince
Albert National Park, Saskatchewan, within the
range of C. |. occidentalis (Goldman 1944) or C. /.
Aquatic Feeding by a Woodchuck!
D. FRASER
NOTES
309
griseoalbus (Hall and Kelson 1959).
Although there is no comprehensive analysis of
specimens throughout North America, the wolves of
the boreal-subalpine forest regions of Alberta and
adjacent areas appear to be the largest of the North
American Gray Wolves.
Literature Cited
Goldman, E. A. 1944. Classification of wolves. Part II. The
wolves of North America. S.P. Young and E.A.
Goldman. American Wildlife Institute, Washington, D.C.
Hall, E. R.and K. R. Kelson. 1959. The mammals of North
America. Volume II. Ronald Press, New York.
Nowak, R.M. 1973. North American Quaternary Canis.
Ph.D. dissertation, University of Kansas, Lawrence,
Kansas.
Skeel, M. A. and L. N. Carbyn. 1977. The morphological
relationship of Gray Wolves (Canis /upus) in national
parks of central Canada. Canadian Journal of Zoology
55(4): 737-747.
Received 2 November 1978
Accepted 19 January 1979
Ontario. Ministry of Natural Resources, Wildlife Research Section, Box 50, Maple, Ontario LOJ 1E0
Fraser, D. 1979. Aquatic feeding by a Woodchuck. Canadian Field-Naturalist 93(3): 309-310.
A Woodchuck (Marmota monax) was seen eating aquatic plants which it obtained by climbing on fallen trees ona lakeshore.
An attraction to sodium-rich plants probably accounts for the behavior.
Key Words: Woodchuck, Marmota monax, aquatic plants, sodium, feeding behavior.
Although they feed on a variety of vegetation,
Woodchucks (Marmota monax) are not generally
known to eat aquatic plants. The following observa-
tions document such behavior.
On 30 June 1978, while canoeing on a small
unnamed lake in Sibley Provincial Park, Ontario
(48°27’N, 88°45’W), I watched through binoculars as
an adult Woodchuck climbed among the tangled
branches of fallen Eastern White Cedar trees (Thuja
occidentalis) overhanging the water. The animal
climbed to a location | to 2 m from shore where it
clung, holding its head 1 to 3 cm above the surface of
the water and its hindquarters slightly higher.
Prominent light areas visible around the nipples
indicated that it was a female. From this location the
Woodchuck dipped its left front paw into the water
and pulled vegetation into its mouth. It fed for about
1 min, then returned to shore, apparently because my
canoe had drifted within 10 m of the location.
The Woodchuck returned to the water’s edge about
8 min later, and walked about | m from shore alonga
half sunken log. There it stopped and chewed on three
plants which it dipped from the water with a front
paw. It then climbed to two sites among the fallen
trees, eating submersed and floating-leafed vegetation
at each location. The animal left to shore 16 min after
it had reappeared.
The feeding sites were dominated by submersed
Hippuris vulgaris and Potamogeton zosteriformis,
floating-leafed Nuphar variegatum, and emergent
Sagittaria sp., growing in 30 cm of water over a soft
organic bottom. At the main feeding location were 10
H. vulgaris plants missing their upper portions, one
uprooted specimen and one damaged fragment of
‘Ontario Ministry of Natural Resources, Wildlife Research Section Contribution No. 78-22.
310
P. zosteriformis, four leaf fragments and two leafless
petioles of N. variegatum, and three damaged
Sagittaria. The second feeding site by the log had
damaged fragments of P. zosteriformis, N. varie-
gatum, and Sagittaria. It appeared that the animal
had completely consumed the portions of H. vulgaris
which it had picked, but left partially chewed
fragments of the other species.
I revisited the site five times in the next 18 days.
On the last day, 18 July, freshly damaged H. vulgaris,
N. variegatum, and Sagittaria were observed. Fresh
damage was not evident on the earlier visits.
Like many herbivores, Woodchucks show evidence
of a sodium-specific hunger in the spring and early
summer (Weeks and Kirkpatrick 1978), and aquatic
plants are generally far richer in sodium than are
terrestrial ones (Jordan et al. 1973). This may explain
why a Woodchuck would make such an effort to
obtain a small quantity of vegetation. Non-aquatic
herbage was readily available on the forest floor,
lakeshore, and roadsides nearby.
Nonetheless, this behavior is clearly a rarity, since
few Woodchucks would be expected to find con-
venient access to aquatic plants in shallow water, and
the animals are rarely seen swimming (Grizzell 1955).
THE CANADIAN FIELD-NATURALIST
Vol. 93
Hamilton (1934) does, however, list one sighting of a
Woodchuck eating Vallisneria americana in July inan
Adirondack Mountain pond. This animal also
reached the vegetation by walking on a fallen log.
I am grateful to D. R. Voigt, M. C. Smith, and
H. P. Weeks for offering suggestions on an earlier
draft of the manuscript.
Literature Cited
Grizzell, R. A. 1955. A study of the southern Woodchuck,
Marmota monax monax. American Midland Naturalist
53(2): 257-293.
Hamilton, W. J. 1934. The life history of the Rufescent
Woodchuck, Marmota monax rufescens, Howell. Annals
of the Carnegie Museum 23: 85-178.
Jordan, P. A., D. B. Botkin, A. S. Dominski, H. S. Lowen-
dorf, and G. E. Belovsky. 1973. Sodium as a critical
nutrient for the moose of Isle Royale. Proceedings of the
North American Moose Conference and Workshop
9: 13-42.
Weeks, H.P. and C.M. Kirkpatrick. 1978. Salt prefer-
ences and sodium drive phenology in Fox Squirrels and
Woodchucks. Journal of Mammalogy 59: 531-542.
Received 20 December 1978
Accepted 12 February 1979
Dispersion of Freshwater Leeches (Hirudinoidea) to
Anticosti Island, Quebec
RONALD W. DAVIES
Department of Biology, University of Calgary, Calgary, Alberta T2N IN4
Davies, Ronald W. 1979. Dispersion of freshwater leeches (Hirudinoidea) to Anticosti Island, Quebec. Canadian Field-
Naturalist 93(3): 310-313.
Four species of freshwater leeches, He/obdella stagnalis, H. triserialis, Glossiphonia complanata, and Mooreobdella fervida
are reported from Anticosti Island. Evidence is present to show that these species dispersed to Anticosti Island passively by sea
currents from Quebec North Shore of the Gulf of St. Lawrence.
Key Words: dispersion, leeches, Hirudinoidea, Anticosti Island, water currents.
Anticosti Island in the Gulf of St. Lawrence is about
216 km long with a maximum width of 48 km. It is
separated by the 29-km or more wide Jacques Cartier
Passage (Détroit de Jacques Cartier) from the North
Shore of the province of Quebec to the north-east, and
by the 64-km or more wide Gaspé Passage (Détroit
d’Honguedo) from the Gaspé Peninsula to the south-
west (Figure |). Having been entirely covered by ice
during the Wisconsin glacial period (Stockwell 1957)
and without known glacial refugia or landbridges to
the mainland during or since the ice recession
(Bleakney 1958; Cameron 1958), the fauna of Anti-
costi Island provides an ideal location to interpret
postglacial migrations and dispersal.
Although Verrill (1863), a prolific author of papers
on leeches (Hirudinoidea), made no reference to
freshwater leeches on Anticosti Island it cannot be
assumed that none were present at that time. Subse-
quently Schmitt (1904) recorded the presence of Dina
(= Mooreobdella) fervida and Johansen (1924) re-
corded Helobdella stagnalis. Ina general review of the
geographical distribution of freshwater leeches of
Canada, Davies (1973) reconfirmed the presence of
M. fervida and H. stagnalis on Anticosti Island and
1979
NOTES 311
a
LAWRENCE
QUEBEC NORTH SHORE
ANTICOSTI ISLAND
st — N ee
NEWFOUNDLAND
50 km
FIGURE |. Map of south-eastern Quebec and Newfoundland showing the major currents (based on Cameron 1958) and the
collection sites of the freshwater leeches (@).
added records for Helobdella triserialis and Glossi-
phonia complanata. Davies (1973) recorded seven
species from the Gaspé Peninsula to which, from
subsequent collections, can now be added Nephelo-
psis obscura and Percymoorensis marmorata (Table
1). Although no new species can be added to the 18
recorded in Quebec by Davies (1973) (Table 1) the
presence on the Quebec North Shore of H. stagnalis,
H. triserialis, G. complanata, M. fervida, N. obscura,
Erpobdella punctata, and Dina parva can be con-
firmed.
Three of the four leech species on Anticosti Island
(H. stagnalis, H. triseralis, and G. complanata) belong
to the family Glossiphoniidae while M. fervida
belongs to the family Erpobdellidae. The Glossi-
phoniidae produce thin-walled cocoons which are
immediately covered by the parents’ body and to
which, while still enclosed within the egg membrane,
the young glossiphoniids develop an embryonic
attachment. As young glossiphonids prematurely
removed from the parent fail to survive, it can be
assumed for these three species at least, that dis-
persion to Anticosti Island must have been in the adult
stage. The Erpobdellidae deposit their cocoons on a
firm substratum and the parent has no further
participation. It is thus possible for M. fervida to be
dispersed in the cocoon stage while attached to stones
or plant material.
TABLE 1—The distribution of freshwater Hirudinoidea
species in the province of Quebec, Gaspé Peninsula,
Anticosti Island, and the Quebec North Shore
Anti- North
Gaspé costi Shore
Que-
Species bec
Piscicola geometra (Linn.)
Piscicola milneri (Verrill)
Piscicola punctata (Verrill)
Bactracobdella picta (Verrill)
Glossiphonia complanata
(Linn.)
Glossiphonia heteroclita
(Linn.)
Helobdella stagnalis (Linn.)
Helobdella triserialis
(Blanchard)
Placobdella ornata (Verrill)
Placobdella phalera (Graf)
Dina parva (Moore)
Erpobdella punctata (Linn.)
Mooreobdella fervida
(Verrill)
Nephelopsis obscura (Verrill)
Bdellerogatis plumbeus
(Moore)
Macrobdella decora (Say)
Mollibdella grandis (Verrill)
Percymoorensis marmorata
(Say)
AA aK MK KK KR KK KK
~*~
AAX~K KK
of
od
3)
The occurrence of four species of freshwater leeches
on Anticosti Island requires explanation, and there
are three possible hypotheses: (a) passive dispersal by
animals, wind, or currents; (b) introduction by man;
(c) active dispersion.
Because of the absence of H. stagnalis from the
Gaspé Peninsula, the leech fauna of Anticosti Island
shows greater similarity to that of the Quebec North
Shore. The absence of a species record from an area is
sometimes a reflection of the degree of collection
intensity, but as there have been more collections from
the Gaspé than from either Anticosti Island or the
Quebec North Shore, the absence of H. stagnalis from
the Gaspé appears to be real. All the leech records
from Anticosti Island are from the north-east Quebec
North Shore-facing coast (Figure |), supporting the
supposition of the Quebec North Shore as the source
of dispersion. Indeed, of the 17 sites sampled on
Anticosti Island (Davies 1973), three of the four sites
on the north-east coast contained leeches but none of
13 on the southern half of the island did.
All four leech species on Anticosti Island are
eurytopic (Davies 1973) benthic feeders, lack jaws,
and are very sensitive to disturbances in the water.
Thus although passive dispersal by birds has been
recorded for benthic leeches (Daborn 1976) it hardly
seems credible that a bird traversing Jacques Cartier
Passage would not occasionally land on the southern
half of this narrow island. Similarly if leeches or
cocoons were transported by strong winds, hurri-
canes, or tornados to Anticosti Island they would be
expected on both halves of the island. As passive
dispersal by wind can probably be discounted,
dispersal by currents or tides remains a possibility.
In a study of the osmoregulatory ability of
freshwater leeches, Reynoldson and Davies (1976)
showed that H. stagnalis was able to maintain itself
hyperosmotically in media between 47 and
112.7 mosmol/L and was a conformer at high-
er media concentrations, and thus could survive
periods of immersion in sea water. Unfortunately the
osmoregulatory abilities of H. triserialis, G. com-
planata, and M. fervida have not been studied but if
they prove to beat least as good as that of H. stagnalis,
it is clear that all four species could survive the sea
passage to Anticosti Island.
Nephelopsis obscura, a widely distributed species in
Canada (Davies 1973), present on the Quebec North
Shore but absent from Anticosti Island can only
maintain itself hyperosmotically between 15.6 and
59.5 mosmol/ L (Reynoldson and Davies 1976) witha
correspondingly low survival rate in salt water.
Ball and Fernando (1970) concluded on the basis of
the dissimilarities between the freshwater triclad
faunae of Anticosti Island, Gaspé, and the Quebec
North Shore, and the similarity on Anticosti Island of
THE CANADIAN FIELD-NATURALIST
Vol. 93
the triclad distribution and amphibian introductions
to the island by man in 1899 (Johansen 1924) that
triclads were also introduced to Anticosti Island by
man. As the triclads and amphibian introductions are
located in the southern half of Anticosti Island,
however, the restriction of leeches to the northern half
“strongly negates the probability of man having
introduced leeches to Anticosti Island.
Active directional dispersion has been recorded for
Percymoorensis marmorata in Quebec (Richardson
1942) and for Erpobdella punctata in Michigan
(Sawyer 1970). But as neither species is recorded from
the Quebec North Shore nor Anticosti Island and
dispersion to Anticosti Island would require an initial
downstream movement into the sea rather than the
upstream movement recorded, active directional
dispersion to Anticosti Island is improbable.
By elimination, the only probable mode of dis-
persion of leeches to Anticosti Island is passive
dispersal by sea currents. For AH. stagnalis, H.
triserialis, and G. complanata the adults could either
be attached to floating debris or free in the water, but
for M. fervida the dispersive place could be either the
cocoons or the adults. The theory of passive dis-
persion of leeches in sea currents to Anticosti Island
from the Quebec North Shore is substantiated by
examination of the predominant currents (Figure 1)
which flow along the Quebec North Shore with a
south-southeast orientation to the mouth of the St.
Lawrence River and then along the Gaspé coast
(Cameron 1958).
It thus appears that the freshwater leeches of Anti-
costi Island have reached the island from the Quebec
North Shore rather than from the Gaspé passively
through the action of sea currents. This is the first
probable record of passive dispersion in sea currents
for freshwater leeches.
The samples from Anticosti Island were collected
by Ian Ball(then of the University of Waterloo) for my
identification.
Literature Cited
Ball, I. R. and C. H. Fernando. 1970. Freshwater triclads
(Platyhelminthes, Turbellaria) from Anticosti Island.
Naturaliste Canadien 97: 331-336.
Bleakney, J.S. 1958. A zoogeographical study of the
amphibians and reptiles of eastern Canada. National
Museum of Canada Bulletin 155: 1-119.
Cameron, A. W. 1958. Mammals of the islands in the Gulf
of St. Lawrence. National Museum of Canada Bulletin
154: 1-165.
Daborn, G. R. 1976. Colonization of isolated aquatic habi-
tats. Canadian Field-Naturalist 90: 56-57.
Davies, R. W. 1973. The geographic distribution of fresh-
water Hirudinoidea in Canada. Canadian Journal of
Zoology 51: 531-545.
1979
Johansen, F. 1924. A biological excursion to Anticosti
Island. Canadian Field-Naturalist 38: 161-164.
Reynoldson, T. B. and R. W. Davies. 1976. A comparative
study of the osmoregulatory ability of three species of
leech (Hirudinoidea) and its relationship to their distribu-
tion in Alberta. Canadian Journal of Zoology 54:
1908-1911.
Richardson, L.R. 1942. Observations on migratory be-
haviour of leeches. Canadian Field-Naturalist 56: 67-70.
Sawyer, R.T. 1970. Observations on the natural history
and behavior of Erpobdella punctata (Leidy) (Annelida:
NOTES
Sis
Hirudinea). American Midland Naturalist 83: 65-80.
Schmitt, J. 1904. Monographie de I’Ile d’Anticosti. Li-
brairie scientifique Paris, A. Hermann, 370 pp.
Stockwell, C. H. 1957. Geology and economic minerals of
Canada. Economic Geology Series 1: 1-517.
Verrill, A. E. 1863. Notes on the natural history of Anti-
costi Island. Proceedings of the Boston Society of Natural
History 9: 132-135.
Received 5 December 1978
Accepted 14 February 1979
Relative Efficiencies of Museum Special, Victor, and Holdfast Traps
for Sampling Small Mammal Populations
ARTHUR M. MARTELL
Canadian Wildlife Service, Great Lakes Forest Research Centre, Sault Ste. Marie, Ontario P6A 5M7
Present Address: Canadian Wildlife Service, 204 Range Road, Whitehorse, Yukon Territory YIA 4Y4
Martell, Arthur M. 1979. Relative efficiencies of Museum Special, Victor, and Holdfast traps for sampling small mammal
populations. Canadian Field-Naturalist 93(3): 313-315.
Small mammal populations in northern Ontario boreal forest were sampled with paired traps. Relative to Museum Specials,
Victors captured proportionally fewer of most species, but Holdfasts captured more soricids, an equal number of cricetids,
and fewer zapodids and sciurids. The most important factors determining the capture rate appeared to be the type of bait
treadle in Museum Special-Victor pairs and trap size in Museum Special—Holdfast pairs. Significant differences in treadle
sensitivity were not reflected in the catch.
Key Words: trap selection, trapping bias, small mammals.
When studying the demography of small mammal
populations it is necessary to obtain relatively
unbiased samples. This is often attempted with one, or
more, of the several types of kill traps commercially
available. Some types have been shown to be
significantly more effective than others in capturing
some species of small mammals (Pruitt and Lucier
1958; Neal and Cock 1969; Smith et al. 1971; Wiener
and Smith 1972). Also the size of the trap has been
shown to influence the mean weight of the catch (Neal
and Cock 1969). Because body weight is often used as
an estimate of age in small mammal studies, estimates
of age structure of the population would also be
biased. Therefore, the type of trap used may sig-
nificantly influence the estimates of the abundance of
the population, the age (weight) structure of the
population, and the relative abundances of the
various species in the community. The purpose of the
present study was to determine which trap-related
biases were present in the small mammal-forestry
research being conducted by the Canadian Wildlife
Service, Ontario Region.
Materials and Methods
Two extensively used small mammal kill traps are
the Museum Special and the Victor mouse trap. The
former measures 69 X 141 mm and has a 12 X 26 mm
wood-covered bait treadle, while the latter measures .
48X99 mm and has an I! X15 mm metal bait
treadle. A third, occasionally used type is the Holdfast
trap, which is the same size as the Victor but has the
same triggering mechanism and treadle as the Mu-
seum Special. All three traps are manufactured by
Woodstream Corporation, Niagara Falls, Ontario
L2E 673, or Lititz, Pennsylvania 17543.
I compared the relative efficiencies of these three
traps by sampling with a Museum Special and either a
Victor or a Holdfast at each trapping point. Strings
were attached to the traps so that they could be placed
in likely capture locations within a |-m radius of the
trapping points, which were approximately 15 m
apart and located either in a single line or in a grid.
Any line or grid contained only one combination of
traps. Traps were baited with a mixture of peanut
butter, rolled oats, and rendered bacon fat, and were
314
checked on 3 consecutive days.
Trapping was conducted in 1976 and 1977 on a
variety of uncut and clearcut boreal forest sites near
Manitouwadge, Ontario (May to October), and ona
series of uncut and strip-cut boreal forest sites near
Beardmore, Ontario (September). A total of 12 324
trapnights was accumulated using Museum Special—
Victor pairs and a total of 20 572 trapnights using
Museum Special-Holdfast pairs (1 trap set for |
night = | trapnight).
The sensitivity of the trap types was measured by
determining the maximum weight, to the nearest
gram, which could be placed on the distal part of the
treadle and still allow the trap to be set. Fifty of each
type of trap were tested after they had been used in the
field for at least four rounds of normal trapping. The
weights used were standard balance weights.
Results and Discussion
The average (x + SE) weight that could be carried
on the distal end of the treadle without tripping the
trap was 1.8+0.08g for Museum _ Special,
3.6 + 0.18 g for Holdfast, and 7.0 + 0.32 g for Victor
traps, which reflects the relative sensitivities of the
traps. The Holdfast held significantly (P<0.001)
more weight than did the Museum Special, and the
Victor held significantly (P<0.001) more weight than
the Holdfast. The range in weights carried was also
different between trap types, with the Victor showing
the greatest range: Museum Special, 1-3 g; Holdfast,
DOs NICOIn Sal Ole:
For Museum Special-Victor pairs I compared the
proportion of Victor-caught animals weighing less
than 10 g, the maximum observed weight carried by a
Victor, with the proportion weighing more than 10 g
and found no significant differences (chi-square test)
in those proportions for soricids (P > 0.1), cricetids
and zapodids combined (P > 0.9), or all three groups
combined (P > 0.1). For Museum Special-Holdfast
pairs I compared the proportion of Holdfast-caught
animals weighing less than 6 g, the maximum obser-
ved weight carried by a Holdfast, with the proportion
weighing more than 6g and found no significant
difference in that proportion for soricids (P > 0.1),
but found that the difference was significant for all
three groups combined (P< 0.001). (All cricetids and
zapodids captured weighed more than 6g.) The
Holdfasts therefore caught a greater proportion of
animals weighing less than 6 g than did the Museum
Specials. The significant differences in triggering
weight were not reflected in the catch.
If triggering weight was not a factor, then the
captures with Museum Special—Holdfast pairs would
be influenced only by trap size, and captures with
Museum Special-Victor pairs would be influenced by
trap size and treadle composition. Table 1 shows that
THE CANADIAN FIELD-NATURALIST
Vol. 93
relative to Museum Specials, Holdfasts caught sig-
nificantly more Masked Shrews (Sorex cinereus) and
total soricids, and significantly fewer Meadow Jump-
ing Mice (Zapus hudsonius) and Least Chipmunks
(Eutamias minimus). Relative to Museum Specials,
Victors caught significantly fewer Masked Shrews,
total soricids, Deer Mice (Peromyscus maniculatus),
Southern Red-backed Voles (Clethrionomys gap-
peri), Heather Voles (Phenacomys intermedius),
Rock Voles (Microtus chrotorrhinus), total cricetids,
and total small mammals. Eight of the nine other
species showed a lower capture rate in Victors than in
Museum Specials.
Trap size appears to be an important factor in
determining the capture rate of three species in
Museum Special—Holdfast pairs. Body size in relation
to trap size may explain the high capture rates of
Masked Shrews in Holdfasts, and Least Chipmunks
in Museum Specials. If so, then treadle composition
clearly outweighs trap size as a factor influencing the
capture rate of Masked Shrews in Museum Special—
Victor pairs.
Smith et al. (1971) and Wiener and Smith (1972)
observed a similar difference in capture rate between
Museum Specials and Victors, and suggested that
Museum Specials were more efficient than Victors
because of their more sensitive triggering mechanism.
My data do not support that suggestion; rather, the
type of bait treadle is indicated as the main factor
influencing the difference in capture rate between
Victors and Museum Specials. The influence may be
due, at least in part, to the wood-covered bait treadle
of the Museum Special which likely retains odors
better than the metal Victor treadle.
There were no significant differences between trap
type in mean body weight of any species captured in
Museum Special—Holdfast pairs (Table 1). In Mu-
seum Special-Victor pairs, however, Southern Red-
backed Voles and Meadow Jumping Mice caught in
Museum Specials were significantly heavier than
those caught in Victors. Differences in mean weight
would be expected to be due to differences in
triggering weight, but that does not appear to be the
cause in my study.
There was no significant difference (chi-square test)
in the proportions of species captured 10 or more
times in Museum Specials and Victors (P > 0.1), but
there was a significant difference between Museum
Specials and Holdfasts (P< 0.001). That difference
was not apparent within either soricids (P > 0.9) or
cricetids (P > 0.1), suggesting that the difference was
between taxonomic groups. When the proportions of
total captures of soricids, cricetids, zapodids, and
sciurids were compared, there was a significant
difference (chi-square test) between Museum Specials
and Holdfasts (P< 0.001) but not between Museum
1979
NOTES
315
TABLE !—Numbers and mean weights (g) of small mammals trapped in Museum Special-Victor pairs (M—V) and Museum
Special-Holdfast pairs (M-H), and the significance (chi-square test and t-test) of the differences within pairs. Sample sizes for
weights approximate those given in the number columns, but in some cases are slightly smaller
Museum Special-Victor
Museum Special—Holdfast
Species Number Weight Number Weight
M Vv M Vv M H M H
Sorex cinereus fal 4|** 355) 3.4 88 160*** 3.5 3.6
Sorex palustris 2 0 15.8 0 1 14.1
Sorex arcticus 2. fed 7.6 3 | 7.1 10.2
Microsorex hoyi 0 l 3.3 0 3 4.1
Blarina brevicauda 7 2 18.5 18.0 8 16 19.9 18.6
Total soricid captures 81 46** 99 Sees
Peromyscus maniculatus 176 67 Ett 15.4 15.4 267 235 16.2 15.8
Clethrionomys gapperi 240 142*** 20.8 19.6* 196 196 Me3) 20.4
Phenacomys intermedius 28 10** 21.0 24.5 36 DF 19.7 23.2
Synaptomys cooperi 2 0 24.4 l 3 24.3 19.9
Microtus pennsylvanicus 14 1} 24.4 23.6 17 21 25.4 24.0
Microtus chrotorrhinus 25 12* 23.5 22.4 6 11 24.2 Des)
Total cricetid captures 485 242*** 523 493
Zapus hudsonius 6 2 ae he 5 0* 14.9
Napaeozapus insignis 2 21.4 0 2 28.3
Total zapodid captures 8 2 5 2
Tamias striatus 0 0 2 2 68.1 91.8
Eutamias minimus 9 7 41.0 45.1 40 hoe? 49.3 47.4
Total sciurid captures 9 7 42 20-%
Total, all species 583 DOU SAX 669 696
*P< 0.05.
**#P< 0.01.
*** P< (0.001.
Specials and Victors (P > 0.5). Relative to Museum
Specials, therefore, Victors captured proportionally
fewer of all species, but Holdfasts captured more
soricids, an equal number of cricetids, and fewer
zapodids and sciurids.
In contrast to my results, other studies have shown
significant differences among species in their capture
rate by Museum Specials and Victors. I tested data
presented by Smith et al. (1971) and Wiener and
Smith (1972) for differences in the proportions of
species captured 10 or more times in Museum
Special-Victor pairs, and found that the difference
was significant in both studies (P< 0.001). Small-
mammal trappers, therefore, should be aware of these
differences and choose their trap type(s) accordingly.
Acknowledgments
I am indebted to the Great Lakes Forest Research
Centre, the Ontario Ministry of Natural Resources,
the Ontario Paper Company and Domtar Ltd. for
their cooperation. I thank A. L. Macaulay for her
assistance in all aspects of the study, and D. A. Welsh,
T. C. Dauphine, J. E. Bryant, and J. F. T. Carreiro
for their comments on the manuscript. =
Literature Cited
Neal, BR. and A.G. Cock. 1969. An analysis of the
selection of small African mammals by two break-back
traps. Journal of Zoology 158: 335-340.
Pruitt, W. O., Jr. and ¢. V. Lucier. 1958. On the relative
efficiency of two kinds of traps. Journal of Mammalogy
39: 157.
Smith, G. C., D. W. Kaufman, R. M. Jones, J. B. Gentry,
and M. H. Smith. 1971. The relative effectiveness of two !
types of snap traps. Acta Theriologica 16: 284-288.
Wiener, J. G.and M. H. Smith. 1972. Relative efficiencies
of four small mammal traps. Journal of Mammalogy 53:
868-873.
Received 30 November 1978
Accepted 5 March 1979
316
THE CANADIAN FIELD-NATURALIST
Vol. 93
Probable Hybrids of Cinnamon X Blue-winged Teal from Southern
Alberta’
D. V. WESELOH and LINDA MCKEANE WESELOH
Provincial Museum of Alberta, 12845 102 Avenue, Edmonton, Alberta T5N 0M6
Present Address: Canadian Wildlife Service, Canada Centre for Inland Waters, P.O. Box 5050, Burlington,
Ontario L7R 4A6
Weseloh, D. V. and Linda McKeane Weseloh. 1979. Probable hybrids of Cinnamon X Blue-winged Teal from southern
Alberta. Canadian Field-Naturalist 93(3): 316-317.
Plumage characteristics of three recent and nine previously reported probable hybrid Cinnamon X Blue-winged Teal are
summarized: males are the rufous color of Cinnamon Teal but retain the white facial crescent and often the white flank patch
of the Blue-winged Teal.
Key Words: Alberta, hybrids, Cinnamon Teal, Blue-winged Teal, Anas cyanoptera, Anas discors.
Until recently most of our knowledge on the
occurrence of Cinnamon (Anas cyanoptera) X Blue-
winged (A. discors) Teal hybridization has come from
aviaries. Kortwright (1942: p. 212) states that“...
even in captivity crosses between the two species are
unknown.” Delacour and Mayr(1945), however, state
that the two species freely interbreed. Spencer (1953:
p. 20) states that “although extremely uncommon
among wild Cinnamon Teal, hybrids are by no means
unknown.” Of the Blue-winged Teal, Delacour (1956;
p. 170) remarks that “In captivity they cross too
readily with Cinnamon Teal, and when both species
are kept on the same pond it is usual to rear a majority
of hybrids . . . This is the more surprising that these
two closely allied Teal seldom cross in the wild state in
the rather narrow areas where they co-exist in North
America.” He goes on to say that Cinnamon Teal
“|. . persecute the weaker Blue-winged Teal...”
In recent years there has been an increasing
number of reports of wild Cinnamon X Blue-winged
Teal hybrids; many of these have come from southern
Alberta and Saskatchewan (Wedgewood and Wedge-
wood 1975; Lang 1973; Lahrman 1971). Also there
have been reports of male Cinnamon Teal and Blue-
winged Teal simultaneously courting a single female
of one of the species (Butot 1974; Andersonand Miller
1953).
In the spring of 1974, we observed three ducks
which we believe were male hybrids of Cinnamon
and Blue-winged Teal. One individual was observed
on 20 May at 16:55, and another at 17:02 at a
slough 0.5 km east of the southeast corner of Eagle
Lake, near Namaka, approximately 56 km east-
southeast of Calgary, Alberta. The first duck was
'Natural History Contribution No. 40, Provincial Museum
of Alberta, Edmonton, Alberta
alone and swam from view within 2-3 min of being
sighted. The second bird was in the company of
two normally plumaged male Cinnamon Teal and a
female typical of the nearly identically appearing
Blue-winged and Cinnamon Teal females. The third
individual was observed by us on 9 June at 19:45,
at Second Vermilion Lake, Banff, Alberta. It was
in the company of two normally plumaged male
Cinnamon Teal and a female of either teal species.
A search of literature as well as present sightings
yields 11 field descriptions of apparently wild Cin-
namon X Blue-winged Teal hybrids. It is not sur-
prising that all individuals have been male as hybrid
females would be virtually undistinguishable in the
field. We have described the major plumage char-
acteristics in Table 1.
In summary, the plumage of male Cin-
namon X Blue-winged Teal hybrids may be char-
acterized as follows: a body having the rufous
coloration of the Cinnamon Teal and the white facial
crescent of the Blue-winged Teal; the white flank
patch of the latter is often present and the breast is
sometimes spotted.
We express our appreciation to Bob Brown,
Michael Bradstreet, and two anonymous reviewers
who made comments on earlier versions of this
manuscript.
Literature Cited
Anderson, W. and A. W. Miller. 1953. Hybridization of
the Cinnamon and Blue-winged Teal in Northern Cali-
fornia. Condor 55: 152-153.
Bent, A.C. 1923. Life histories of North American
wildfowl. Part 1. U.S. National Museum Bulletin 126.
Reprinted 1951.
Butot, R. 1974. Ménage a trois, or the end of a species.
Calgary Field Naturalist 5(11): 281-282.
1979
NOTES
SHl7/
TABLE 1—Plumage features of male Cinnamon X Blue-winged Teal hybrids
White facial White flank Breast
Body coloration crescent patch spotting Location and source
1. Dark cinnamon Distinctive Distinctive Utah; Wilson and Van den Akker 1948
2. Cinnamon Snow-white Colorado; Bent 1923
3. Cinnamon Distinct Believed lacking Utah; Spencer 1953
4. Cinnamon Prominent Alberta; Spencer 1953
5. Cinnamon Clear Indistinct California; Anderson and Miller 1953
6. Reddish Washed out* Prominent* Present Saskatchewan; Lahrman 1971
7. Cinnamon Faint Alberta; Lang 1973
8. Rusty red Small Light Saskatchewan; Wedgewood and Wedgewood 1975
9. Cinnamon Prominent Prominent Alberta; this paper
10. Cinnamon Washed out Absent Alberta; this paper
11. Rufous-cinnamon Washed out Washed out Alberta; this paper
*Authors’ interpretation from photo.
Delacour, J. 1956. The waterfowl of the world. Country
Life Ltd., London. Volume 2. 232 pp.
Delacour, J. and E. Mayr. 1945. The Family Anatidae.
Wilson Bulletin 57: 3-55.
Kortwright, F. H. 1942. Ducks, geese and swans of North
America. Stackpole Company, Harrisburg, Pennsy]l-
vania and Wildlife Management Institute, Washington.
llth printing. 476 pp.
Lahrman, F.W. 1971. Hybrid Cinnamon Teal X Blue-
winged Teal at Regina. Blue Jay 29: 28.
Lang, V. 1973. Observation of probable Cinnamon Teal
(Anas cyanoptera) X Blue-winged Teal (Anas discors)
hybrid. Calgary Field Naturalist 5(4): 139.
Spencer, H.E., Jr. 1953. The Cinnamon Teal (Anas
cyanoptera Vieillot): Its life history, ecology and man-
agement. M.Sc. thesis, Utah State Agricultural College,
Logan, Utah. 184 pp.
Wedgwood, S.L. and J. A. Wedgwood. 1975. Probable
Cinnamon Teal — Blue-winged Teal cross. Blue Jay
33(3): 181-182.
Wilson, V.T. and J.B. Van den Akker. 1948. A hybrid
Cinnamon Teal - Blue-winged Teal at the Bear River
Migratory Bird Refuge, Utah. Auk 65: 316.
Received 20 October 1976
Resubmitted 28 February 1979
Accepted 5 March 1979
Common Garter Snake Predation on Ring-billed Gull Chicks
PETER M. FETTEROLF
Department of Zoology, University of Toronto, Toronto, Ontario MSS I1A1
Fetterolf, P.M. 1979. Common Garter Snake predation on Ring-billed Gull chicks. Canadian Field-Naturalist
93(3): 317-318.
A Common Garter Snake was observed eating two freshly hatched Ring-billed Gull chicks.
Key Words: Common Garter Snake, Ring-billed Gull chicks, predation, Thamnophis sirtalis, Larus delawarensis.
In this note I describe an act of predation upon
Ring-billed Gull (Larus delawarensis) chicks by a
Common Garter Snake, Thamnophis sirtalis. The gull
colony is on Mugg’s Island, Toronto Harbor,
Toronto, Ontario. The observation was made froma
blind situated in a Ring-billed Gull colony of
approximately 6000 pairs. About 50 pairs of Herring
Gulls, L. argentatus, also nest on the island.
The incident began at 07:35 EST on 25 May 1978 as
the snake moved from beneath the platform which
supports the blind. The 80-cm reptile moved approxi-
mately 4 m to the edge of a gull nest which contained
three chicks. Two chicks were about 48 h old and the
third was less than 24 h old. Freshly hatched chicks
average 35-45 g. The snake tested one half of an
eggshell lying at the nest rim with its tongue, grasped
the shell between its jaws, and moved it about 20 cm
from the nest. The snake then raised its head
approximately Scm above the substrate, froze
momentarily, and lunged rapidly at a chick, seizing it
by the bill. The snake engulfed the chick up to the
wings within 4 min and then writhed several times.
318
The chick was gone from view 14 min after capture,
and the snake remained stationary for a few minutes
more. As it contorted slowly and rubbed its nose in the
sand, the obvious bulge moved posteriorly.
Following this respite, the reptile inched its way
toward a nest about | mdistant which contained eggs.
The contents were tested with the tongue, and the
snake continued to two adjacent nests with eggs where
it performed the same operation. Finally, a nest with
two 48-h-old chicks was reached. The two chicks were
different in size, and the Garter Snake seized the
smaller one by the throat. The snake released the chick
long enough to grasp it by the bill as it had done with
the first victim. The swallowing behavior which
followed was similar to that in the first predatory act
23 min earlier, and |! minafter it grasped the chick the
snake disappeared beneath the blind. At that time,
80% of the chick had disappeared within the snake’s
jaws.
The gulls’ reaction to the snake was interesting.
Hatching had just started in the area and during the
entire incident all chicks remained in the nest bowls or
on their rims. Adult gulls stood about 0.75 m from the
snake and called loudly, using the “Kow-Kow’ call
described by Moynihan (1956). Adults displayed no
anti-predator behavior (Krunk 1964) and none took
flight at the sight of the reptile.
Gull chicks are consumed by avian and mammalian
predators such as the Great Horned Owl, Bubo
virginianus (Burger 1974); Marsh Hawk, Circus
cyaneus (Burger 1974); Mink, Mustela vision (Burger
1974); Hedgehog, Erinaceus europaeus (Kruuk 1964);
Red Fox, Vulpes vulpes (Kruuk 1964; Burger 1974).
Campbell (1969) reported that the Wandering Garter
Snake, 7. elegans vagrans, preyed upon Glaucous-
winged Gull, L. glaucescens, chicks. Birds are rare in
the diet of 7. sirtalis (Hamilton 1951; Fouquette 1954;
Fitch 1965; Gregory 1978). Thus, consumption of
Ring-billed Gull chicks may be a special case of
opportunistic feeding. Campbell’s observations of
Wandering Garter Snake predation on Glaucous-
winged Gull chicks occurred in an ecological situation
similar to that on Mugg’s Island. In both cases, the
snakes were the only observed reptilian inhabitants of
the islands used by nesting gulls. Perhaps the snakes,
which use olfaction to detect food (Fox 1952;
Burghardt 1970), were originally attracted to the
colonies by the odor of fish scraps dropped by the
gulls. The Wandering Garter Snake ate food pellets
regurgitated by gulls (Campbell 1969). The fact that
THE CANADIAN FIELD-NATURALIST
Vol. 93
the snake on Mugg’s Island struck at and grasped the
fresh eggshell, which was unlikely to smell of fish,
suggests that the snake may have utilized chick odor in
its hunting behavior. It is difficult to assess the
frequency of gull chick predation by the Common
Garter Snake on Mugg’s Island, for it has only been
observed once in over 1200 h of observations in the
past 4 yr. Therefore the incident reported here is
probably an example of a predator taking a locally
abundant, easily obtained food source not normally in
its diet.
Acknowledgments
My thanks go to D. W. Dunham who provided
financial support from a National Research Council
of Canada operating grant, to A. R. Gibson who
provided some garter snake references, and to my wife
who did the typing of the manuscript.
Literature Cited
Burger, J. 1974. Breeding adaptations of Franklin’s Gull
(Larus pipixcan) to a marsh habitat. Animal Behavior
22: 521-567.
Burghardt, G. M. 1970. Intraspecific geographical varia-
tions in the chemical food cue preference of newborn
garter snakes. Behavior 36: 246-257.
Campbell, R. W. 1969. Notes onsome foods of the wander-
ing garter snake on Mitlenatch Island, British Columbia.
Syesis 2: 183-187.
Fitch, H. S. 1965. An ecological study of the garter snake,
Thamnophis sirtalis. University of Kansas Publications,
Museum of Natural History 15: 393-464.
Fouquette, M. J., Jr. 1954. Food competition among four
sympatric species of garter snakes. Texas Journal of
Science 1964 (2): 172-188.
Fox, W. 1952. Notes on the feeding habits of Pacific
coast garter snakes. Herpetologica 8: 4-8.
Gregory, P. J. 1978. Feeding habits and diet overlap of
three species of garter snakes (Thamnophis) on Vancouver
Island. Canadian Journal of Zoology 56(9): 1967-1974.
Hamilton, W. J., Jr. 1951. The food and feeding behavior
of the garter snake in New York state. American
Midland Naturalist 46: 385-390.
Kruuk, H. 1964. Predators and anti-predator behavior of
the Black-headed Gull (Larus ridibundus L.). Behavior
Supplement I1.
Moynihan, M. 1956. Notes on the behavior of some North
American gulls. 1. Aerial hostile behavior. Behavior 10:
126-179.
Received 3 November 1978
Accepted 3 February 1979
1979
NOTES
319
The Rock Vole (Microtus chrotorrhinus) as a Transition Zone Species
GORDON L. KIRKLAND, Jr. and CHARLES M. KNIPE
Vertebrate Museum, Shippensburg State College, Shippensburg, Pennsylvania 17257
Kirkland, Gordon L., Jr. and Charles M. Knipe. 1979. The Rock Vole (Microtus chrotorrhinus) as a Transition zone
species. Canadian Field-Naturalist 93(3): 319-321.
Rock Voles (Microtus chrotorrhinus) were collected in mixed northern hardwood forests in New York’s Adirondack
Mountains at elevations between 460 and 700 m. These voles were much more common in the Transition zone habitats than
previously thought. Data are presented on the habitat, relative abundance, and ecological associates of the 43 specimens.
Key Words: Rock Vole, Microtus chrotorrhinus, Adirondack Mountains, Canadian zone, Transition zone.
Rock Voles (Microtus chrotorrhinus) have tradi-
tionally been considered to be boreal small mammals.
Recent research at the periphery of their range
indicates that Rock Voles are not only geographically
more widespread than previously believed, but they
have greater ecological amplitude as well (Buech et al.
1977; Kirkland 1977; Roscoe and Majka 1976; Timm
1974: Timm et al. 1977). In the northeastern United
States, Rock Voles have been thought to be limited in
distribution to higher elevations and more boreal
habitats. Martin (1971a) states that the Rock Vole is
most closely associated with the Canadian life zone.
Furthermore, he notes that records of the Rock Vole
in the Transition zone may be explained on the basis
of expansion of the Rock Vole’s distribution under
suitable conditions from the sanctuary of more
favorable Canadian zone habitat. Examination of the
distribution of captures of the Rock Vole in New
York’s Adirondack Mountains prior to 1970 tends to
confirm this traditional view of the ecological distri-
bution of the Rock Vole. With the exception of a
series of 29 specimens collected by C. F. Batchelder at
Keene Heights (elevation 400 m), 61 of 91 specimens
examined in museum collections were taken at
elevations above 976 m, which is the lower limit of
Red Spruce (Picea rubens)-dominated forests in the
Adirondacks (Braun 1964). Since 1970, however, 43
Adirondack Rock Voles collected by Shippensburg
State College Vertebrate Museum field personnel
have been trapped at elevations below 1100 m. In
addition, F. J. Jannett, Jr. (Cornel! University) and
R. Rosen (University of Vermont) (personal com-
munications) have obtained for use in laboratory
studies 20 live Rock Voles at elevations between
640 and 760m in the Adirondacks. These recent
records reveal that Rock Voles are more widespread
and abundant in Transition zone habitats than was
previously believed. This paper presents information
on the habitat, relative abundance, and ecological
associates of 43 M. chrotorrhinus captured from
1973-1978 at elevations of 460 to 700 m in Essex
County, New York.
Methods
The 43 Rock Voles described in the paper were
collected in 10 traplines (Table 1) representing seven
localities, as follows: New York, Essex County, St.
Hubert’s, 2.7 km SE, elevation 457 m (1); Tahawus,
0.40 km SE, elevation 549 m (1); 1.9 km W, elevation
549m (1); 7.9km NNW, elevation 640m (26);
8.7 km N, 3.2 km W, elevation 670 m (9); Wallface
Mountain, 0.8km SW, elevation 700m (4);
3.2 km SW, elevation 625 m (1). The specimens were
obtained as part of a small mammal survey of Essex
County, New York. Sampling procedures involved
establishing traplines with variable numbers of sta-
tions, usually with three snap-traps per station. Mu-
seum Specials baited with rolled oats were used princi-
pally, although each line might have up to 10% rat and/
or mouse traps. At each station, the traps were set with-
in 2 m of each other, and traplines were operated for
either 2 or 3 d. The 10 traplines yielding Rock Voles
had a total sampling effort of 2691 trapnights (TN).
Results and Discussion
The trapline and specimen records for the 43 Rock
Voles reveal that 42 were captured in typical Transi-
tion zone forests dominated by Yellow Birch (Betula |
lutea), Sugar Maple (Acer saccharum), and American
Beech ( Fagus grandifolia). The only exception was the
single capture along a stream, 2.7 km SE St. Hubert’s,
in a Red Spruce — Eastern Hemlock ( Tsuga canaden-
sis) association. All specimens were captured at sites
where rocks or boulders were conspicuous habitat
components and the canopy was semi-open. Six of the
seven localities at which 42 of the Rock Voles were
captured had streams of varying sizes flowing through
them. Martin (1971a) notes that 47 of 83 Rock Voles
he collected were caught within 30 ft (9 m) of running
water, either surface or subsurface.
The microhabitats of the Rock Voles in this study
tended to be characterized by the presence of rocks,
mosses, ferns, and forbs (e.g., Oxalis, Clintonia,
Viola, and Smilacina). The percentages of Rock Voles
collected within 1m of these four microhabitat
320
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 1—Small mammals captured in 10 traplines yielding Rock Voles in Transition zone habitats of Essex County, New York
Trapline number, year, elevation (m), sampling effort (TN)
Ie 2
4;
73, NOS.
549, 457,
Species 450 44]
Masked Shrew (Sorex cinereus)
Long-tailed Shrew (Sorex dispar) 2
Smokey Shrew (Sorex fumeus) 3)
Water Shrew (Sorex palustris)
Short-tailed Shrew (Blarina brevicauda) 5 4
Chipmunk (Tamias striatus) 2
Red Squirrel (Tamiasciurus hudsonicus)
Northern Flying Squirrel (Glaucomys sabrinus)
Deer Mouse (Peromyscus maniculatus) 6 8
Red-backed Vole (Clethrionomys gapperi) ] 16
Rock Vole (Microtus chrotorrhinus) I I
Meadow Vole (Microtus pennsylvanicus)
Woodland Jumping Mouse ( Napaeozapus insignis) 3} I
Meadow Jumping Mouse (Zapus hudsonius)
Totals 18 35
a 4, 5, 6. i 8, 9, 10, Totals
1975, P1975," 1975, 1977, 1977, 1978, ealo7sammnenee
549, 625, 700, 640, 640, 640, 640, 700,
540 300 300 240 240 120 120 240 2691TN
6 3 I 2 Ai, LEG
I 2 I 2 I 9
2 I 6
2 2
24 12 5 7 10 I 1 69
3 6
I I
I 2
16 14 il 4 2 2 2 4 69
14 8 in uae 825 84
I 4 14 10 I Omens
I I
1 4 5 4 2 2 2 34
I I
[Sie merase 143 OES 9 9 19 343
components were as follows: rocks (66%), mosses
(88%), ferns (85%), and forbs (91%). Live ground cover
at individual capture sites varied from less than 25 to
more than 75% but averaged 50-60%. The Rock Voles
appeared to spend a considerable portion of their time
in subterranean activity. In this survey, 53.5% of the
specimens were caught in traps set under rocks or
below the surface of rocky sites, and 65% of the total
were caught in “unexposed” sites, either under rocks,
logs, roots, or overhangs.
The 43 Rock Voles were captured in association
with 13 other species of small mammals (Table 1).
They were most frequently captured in association
with Deer Mice (Peromyscus maniculatus gracilis),
Short-tailed Shrews ( Blarina brevicauda), and Wood-
land Jumping Mice (Napaeozapus insignis).
Although collected in only 7 of 10 traplines, Red-
backed Voles (Clethrionomys gapperi) were the most
abundant ecological associate with a total of 84
specimens. Martin (1971b) noted that he collected a
“relatively unchanging proportion of approximately
three C. gapperi for every M. chrotorrhinus in
favorable rock vole habitat.” In this study, the
proportion of C. gapperi to M. chrotorrhinus was
2.0:1.0; however, this varied considerably between
traplines. In individual traplines, the ratio of these
two species ranged from 16:1 in favor of C. gapperito
9:0 in favor of M. chrotorrhinus.
In October of 1965 and 1966, J. N. Layne and
students from Cornell University sampled small
mammals on Whiteface Mountain, Essex County,
New York. They collected 20 M. chrotorrhinus at
elevations from 1128 to 1433 min 10 of their traplines,
with a total sampling effort of 1069 TN. These previ-
ously unpublished data permit some comparisons of
Rock Voles inhabiting Canadian and Transition
zones in the Adirondacks. The catch per unit sampling
effort of 1.87/100 TN for the Whiteface Mountain
sampling was slightly higher but did not differ
significantly from the 1.60/ 100 TN for the Transition
zone sampling in this study (0.75 > P > 0.50). This
indicates that in the Adirondacks, Rock Voles may
have comparable population levels in suitable Can-
adian and Transition zone habitats. The 20 Rock
Voles on Whiteface Mountain were captured in
traplines yielding seven other species of small mam-
mals as follows: Masked Shrew (Sorex cinereus) (3),
Long-tailed Shrew (Sorex dispar) (6), Smokey Shrew
(Sorex fumeus) (1), Short-tailed Shrew (3), Deer
Mouse (12), Red-backed Vole (27), and Meadow Vole
(Microtus pennsylvanicus) (5). It is of interest that on
Whiteface Mountain, the ratio of Red-backed Voles
to Rock Voles was only 1.4:1.0. A Spearman Rank
Correlation analysis (Siegel 1956) of the species of
small mammals collected on Whiteface Mountain and
in the Transition zone habitats (Table 1) reveals a
significant correlation (rs = 0.696, P< 0.05) between
the relative abundance of species in small mammal
communities containing Rock Voles in the two
ecological zones. Thus, although the Rock Voles are
occupying dissimilar plant communities, they are
members of qualitatively similar small mammal
communities.
The recent capture of Rock Voles at a variety of
locations in the Transition zone of the Adirondack
Mountains suggests that traditional assessments of
1979
this species’ ecological distribution should be re-
evaluated. Rather than being restricted primarily to
boreal habitats characteristic of the Canadian zone,
the Rock Voles also occupy sites in northern
hardwood forests where appropriate microhabitat
components are present, specifically rocks, flowing
water, mosses, ferns, and forbs.
Acknowledgments
We thank James N. Layne for permitting us to use
data he and his students had collected on Rock Voles
at Whiteface Mountain and for critically reviewing a
draft manuscript. Our thanks are also extended to
Walter M. Chapman, Manager, of the MacIntyre
Development, N L Industries, Tahawus, New York
for providing housing and access to private lands
sampled in this study. We acknowledge Carol J.
Kirkland, Henry W. Setzer, and the students from
Shippensburg State College who assisted in collecting
the specimens described in this paper.
Literature Cited
Braun, E. L. 1964. Deciduous forests of eastern North
America. Hafner Publishing Company, New York. 596pp.
Buech, R. R., R. M. Timm, and K. Siderts. 1977. Asecond
population of the Rock Vole, Microtus chrotorrhinus, in
NOTES
321
Minnesota with comments on habitat. Canadian Field-
Naturalist 91: 413-414.
Kirkland, G.L., Jr. 1977. The Rock Vole, Microtus
chrotorrhinus (Miller) (Mammalia: Rodentia) in West
Virginia. Annals of the Carnegie Museum of Natural
History 46: 45-53.
Martin, R. L. 1971a. The natural history and taxonomy of
the Rock Vole, Microtus chrotorrhinus. Ph.D. thesis,
University of Connecticut, Storrs. 123 pp.
Martin, R.L. 1971b. Interspecific associations of Rock
Voles. Beta Kappa Chi Bulletin 30(2): 5-7.
Roscoe, B. and C. Majka. 1976. First record of the Rock
Vole (Microtus chrotorrhinus) and the Gaspé shrew
(Sorex gaspensis) from Nova Scotia and the second record
of Thompson’s pygmy shrew (Microsorex thompsoni)
from Cape Breton Island. Canadian Field-Naturalist
90: 497-498.
Siegel, S. 1956. Nonparametric statistics for the behavioral
sciences. McGraw-Hill Book Company, New York. 312 pp.
Timm, R.M. 1974. Rediscovery of the Rock Vole
(Microtus chrotorrhinus) in Minnesota. Canadian Field-
Naturalist 88: 82.
Timm, R.M., L.R. Heaney, and D.D. Baird. 1977.
Natural history of Rock Voles (Microtus chrotorrhinus) in
Minnesota. Canadian Field-Naturalist 91: 177-181.
Received 21 December 1978.
Accepted 12 March 1979.
Eastern Limit of the five-lined Skink, Ewmeces fasciatus, in Ontario
RICHARD D. USSHER! and FRANCIS R. COOK2
IRR 1, Delta, Ontario KOE 1G0
2Herpetology Section, National Museum of Natural Sciences, Ottawa, Ontario KIA 0M8
Ussher, Richard D. and Francis R. Cook. 1979. Eastern limit of the Five-lined Skink, Eumeces fasciatus, in Ontario.
Canadian Field-Naturalist 93(3): 321-323.
New records are reported for the Five-lined Skink, Eumeces fasciatus, from Frontenac County (Palmerston Lake; 2.4 km W
of Snow Road: and 1.6 km SE of Snow Road Station), Lanark County (Robertson Lake), and Leeds and Grenville (Landon
Bay) County. The similarities between the eastern range limit in Ontario of the skink and those of the Northern Ribbon Snake
and Black Rat Snake are discussed; the adjacent more poorly-drained lowlands are suggested as a distribution barrier.
Based on collections reported by Patch (1934),
Logier and Toner (1961, p. 59) cited Arden and
Mountain Grove in Frontenac County as the most
eastern localities at which the Five-lined Skink,
Eumeces fasciatus, had been taken in Canada.
Records accumulated since this publication allow a
better definition of the eastern limit of its distribu-
tion.
On 30 August 1961, seven skinks were found by
Cook and M. Gordon Foster 1.5 mi (2.4 km*) W of
Highway 509 at Snow Road, Frontenac County (these
are now in the National Museums of Canada,
Collection Number 5618). They were inhabiting a
rocky outcropping in the vicinity of a deserted hillside
farmsite. One individual was discovered under a
board at the farmsite: the others were under generally
flat rocks in the adjacent woods and hillside. The
woods were primarily White Pine (Pinus strobus) but
a mixture of deciduous trees was also present. A thick
layer of pine needles ‘covered most of the ground in the
wooded area. Old lumber piles and scattered loose
boards were prevalent at the farm site, and loose rocks
were common in the woods and on the hillside.
Additional records in this region include a single
322
individual taken by D. Scobie on 31 July 1960 from
near Palmerston Lake, about 2 mi (3.2 km*) up the
lake from Ompah (NMC 5110): one specimen
collected on 10 August 1961 by A. G. Walker | mi
(1.6 km*) SE of Snow Road Station, Palmerston
Township, Frontenac County (Carleton University
Museum of Zoology; D. A. Smith, personal com-
munication); and one specimen collected by F.C.
Zufeldt in May 1962 from Robertson Lake (NE of
Lavant), Lanark County (NMC 6507). The map in
Conant (1975) is based on the National Museum of
Natural Sciences records, but individual localities
have not been previously published.
Luciuk and McCabe (1971) noted that the species is
reported to occur at “3rd Depot Lake” and at Perth
Road Bay near Chaffey’s Locks. The latter locality is
about midway between Snow Road Station and
Gananoque.
On 22 August 1973 at the Landon Bay Campsite on
the north shore of the St. Lawrence River, Ussher
observed a single individual. The locality was a bare
patch of Precambrian rock, some 30 m above the
water of Landon Bay, 8.5 km E of Gananoque in
Leeds and Lansdowne Township, Leeds and Gren-
ville County. This sighting is the only one from the
area by Ussher, although he became a resident of
Leeds and Grenville County in 1970, and visited this
locality and similar ones on many occasions.
East of these records, in the Ottawa District, (a 48-
km radius around the National Capital) where
herpetofaunal surveys have been undertaken by the
National Museum of Natural Sciences for several
decades, and near Bishop’s Mills in Leeds and
Grenville County, where Cook has resided since 1970,
there have been neither sightings nor reports of
skinks.
Of interest is the roughly similar eastern limit of two
other Ontario reptiles, the Black Rat Snake, Elaphe
obsoleta obsoleta, which is known to occur from just
south of Smith’s Falls to Mallorytown (NMNS files
including personal communications from R. V. Lind-
say, John Woods, Roger E. Roy, Harold Parsons, and
Al MacDonald), and the Northern Ribbon Snake,
Thamnophis sauritus septentrionalis, which reaches
the “Horseback Mountains” near Pakenham (Cook
1968) and slightly beyond (Tony Tobias, personal
communication) and Mallorytown Landing (Woods
and Cook 1976). In his review of the herpetofauna of
the Thousand Islands Region, New York, Werner
(1959) reported both the Black Rat Snake and the
Ribbon Snake, but not the Five-lined Skink.
None of these three species have been reported from
east of the rough and relatively well-drained elevated
terrain of the Precambrian Shield topography, onto
*Collectors originally stated distances in whole or half miles.
THE CANADIAN FIELD-NATURALIST
Vol. 93
the adjacent low-lying, and relatively more poorly-
drained area which was covered by the postglacial
Champlain Sea of the Ottawa and St. Lawrence
valleys. Nor have any been reported on the eastern
side of this presumed barrier, in the Precambrian of
the Gatineau and Laurentian regions between Ottawa
and Montreal.
The populations of these three species reported in
Leeds and Grenville, Frontenac, and Lanark counties
may be prevented from colonizing the lowlands at
the northeastern edge of their range by a lack of well-
drained hibernating sites and the lack of sufficient
insulation that is provided by deep accumulations of
snow in the hollows of the adjacent broken top-
ography. The moderating effect of the Rideau Lakes
system on local microclimates, possibly promoting
warm pockets, and the less intensive agriculture on its
rough terrain, leaving more of the area in forest, are
additional factors that may contribute to the survival
of these species in the area. Their eastern limit roughly
corresponds to the eastern boundary of Herpeto-
faunal Section | defined by Bleakney (1958), which is
based on mean July temperature and the length of the
growing season.
A fourth species, the Stinkpot, Sternotherus odor-
atus, reaches its eastern limit in the general area, but it
invades the lowlands along the Rideau and Missis-
sippi rivers at least as far as Beckets Landing (NMC
3938) in Leeds and Grenville counties and Innisville
(NMC 2157) and Pakenham (NMC 13777) in Lanark
County. Perhaps its different overwintering habitat,
in these rivers does not place on it the same constraints
for hibernation sites that are suggested for the terres-
trially hibernating species discussed above.
Additional observations of any of these species at
the eastern edge of their range, giving locality, habitat,
date, number observed, and observer(s) name(s)
would be appreciated by the authors. Information on
the status of the Black Rat Snake is also being
collected for the Ontario Ministry of Natural Re-
sources by Harold Parsons and Al Macdonald as part
of a detailed study of it in the Rideau Lakes region,
and by the Federation of Ontario Naturalists in a
general survey of its Ontario range. Because sightings
of any of these species are often a matter of chance,
and because populations may be scattered and
fluctuating in abundance from year to year, the
cumulative interest and efforts of all observant
naturalists is needed further to define their ranges. A
more precise discussion of these limiting factors must
wait until such observations are available, and should
also include comparison with the northern limit of
these species. The latter may roughly correspond (with
the notable exception of the Black Rat Snake) with
the southern boundary of the 300-m contour and a
large height of land covering the Haliburton and
72)
Algonquin highlands (see map in Weller and Palermo
1976: W.G. Sprules and W.F. Weller, personal
communication).
Literature Cited
Bleakney, J. Sherman. 1958. A zoogeographic study of the
amphibians and reptiles of eastern Canada. National
Museum of Canada Bulletin 155: 1-119.
Conant, Roger. 1975. A field guide to reptiles and am-
phibians of eastern and central North America. Houghton
Mifflin Company, Boston. xvii + 429 pp.
Cook, Francis R. 1968. Lizards and snakes of the Ottawa
District. Trail & Landscape 2(4): 99-106.
Logier, E. B.S. and G. C. Toner. 1961. A check list of the
amphibians and reptiles of Canada and Alaska. Second
Edition. Royal Ontario Museum, Life Sciences Division,
Contribution 53: 1-93.
Barn Owls in Quebec
NORMAND DAVID
NOTES
323
Luciuk, Lubomyr and Jim McCabe. 1971. The Five-Lined
Skink (Eumeces fasciatus). Canadian Herpetologists
Society Quarterly 1(2): 3.
Patch, C. L. 1934. Euwmeces in Canada. Copeia 1934(1):
50-51.
Woods, John G. and Francis R. Cook. 1976. Range
extension of the Ribbon Snake in Eastern Ontario.
Canadian Field-Naturalist 90(1): 69-70.
Werner, William E. 1959. Amphibians and reptiles of the
Thousand Islands area, New York. Copeia 1959(2): 170-
172.
Weller, Wayne F. and R. Victor Palermo. 1976. A northern
range extension for the Western Chorus Frog, Pseu-
dacris triseriata triseriata (Wied), in Ontario. Canadian
Field-Naturalist 90(1): 163-166.
Received 10 November 1978
Accepted 16 March 1979
Centre de recherches écologiques de Montréal, Case postale 6128, Succursale “A,” Montréal, Québec H3C 3J7
David, Normand. 1979. Barn Owls in Quebec. Canadian Field-Naturalist 93(3): 323-324.
An evaluation of Barn Owl records in Quebec shows that there is no conclusive evidence that the species has actually bred
within the province.
Key Words: Tyto alba, Quebec.
Information from F. Créte, then Director of the
Museum of the Deaf and Dumb Institute of Mon-
treal, provided the only suggestion of breeding of the
Barn Owl (Tyto alba) in Quebec, at Berthierville,
Berthier County, in 1931 (Cayouette 1947; Godfrey
1966). The unpublished ornithological notes of the late
Victor Gaboriault, however, show that breeding of
Barn Owls in Berthierville in 1931 was never substan-
tiated. An evaluation of that record and of other
reported occurrences in the province indicates that the
Barn Owl is a casual visitor, but gives no indisputable
evidence that it breeds here.
Early in 1945, R. Cayouette learned that a “dozen
Barn Owls” had been seen in Berthierville around
1931, and he was referred to Créte for further details.
Créte indicated that a pair, discovered by A. Paquette,
had nested in the church steeple, that from the four
young he had received one in the flesh which was
mounted by a local taxidermist, and that Paquette
had given him a mounted specimen on 29 December
1931. Créte added that he kept the second specimen
and gave the first to the museum of College de Sainte-
Anne-de-la-Pocatiére. With his letter Crete included
copies of the label and of the accession card of the
specimen given to him by Paquette. According to the
label the specimen (No. 31/16) is an adult male
captured on 10 November 1931; on the accession card,
however, Créte wrote that this bird was killed in
November 1931, from a brood of four whose parents
had settled in the church steeple.
On 28 March 1946, Gaboriault interviewed Paquette
and was told that high winds had been blowing for
several days when about 20 Barn Owls appeared in the ©
Berthierville Islands in 1931. After a while they took
refuge in the church steeple where Paquette killed
eight birds. He mounted one which is at the Deaf and
Dumb Institute, and sent the others to various
museums. As the steeple floor was covered with
droppings, it was evident several birds had been there
for some time. Paquette could not give more definite
details.
Paquette’s recollection is confirmed by a post-
humous article (Paquette 1961) which he read in
French at the fiftieth meeting of the AOU at Quebec
City in 1932 (see The Auk S50, p. 74). Relating his
observations on birds of prey in captivity he said that
about 15 Barn Owls had settled in Berthierville, and
that he kept one bird for a week in order to study its
postures before mounting it.
Gaboriault’s list of Barn Owl specimens taken in
324
TABLE 1—Records of occurrence of the Barn Owl in Quebec
THE CANADIAN FIELD-NATURALIST
Vol. 93
Locality Date Remarks! References
Longueuil October 1915 Captured by W. J. Low V. Gaboriault
L’Assomption November 1926 Captured Auk 64, p. 631
Longueuil 1 November 1931 Deaf and Dumb Institute Museum V. Gaboriault
November 1931
10 November 1931
15 June 1936
Berthierville
Berthierville
St.Félix-de-Kingsey
Beauharnois 12 May 1944 Captured by V. Gaboriault
Thetford Mines 3 September 1944 Captured in a barn
Saint-Hubert 1961 One “pair” seen in summer
Giffard 18, 24 June 1963 Sight records
14 April 1967
30 May 1968
20 December 1970
22 December 1970
Berthierville Sight record
Mont-Carmel
Cap Tourmente
Cap Tourmente
Found dying
Sight record
Rigaud 12 June 1971 Captured in a barn
Cap Tourmente 8 May 1974 Found dead
Near Montreal Spring 1975 Shot
Huntingdon Fall 1975 Killed by a car
Saint-Coeur-de-Marie 1975 Captured in a barn
Masson July 1977
condition)
Captured by J. Desjardins
At least 3 captured by A. Paquette
Captured in a barn
Captured in a barn
Captured (female in breeding
V. Gaboriault
V. Gaboriault
PSNHC? Annual Report 1938, p. 161
V. Gaboriault
Auk 64, p. 631
PQSPB3 Annual Report 1961, p. 25
Bulletin ornithologique 8(4), p. 3
Bulletin ornithologique 12, p. 19
Bulletin ornithologique 13, p. 37
Bulletin ornithologique 15, p. 95
Bulletin ornithologique 15, p. 95
Bulletin ornithologique 16, p. 58
Bulletin ornithologique 19, p. 24
American Birds 29, p. 830
American Birds 30, p. 696
Bulletin ornithologique 21, p. 54
Henri Ouellet (pers. comm.)
‘One individual only unless otherwise stated.
2The Provancher Society of Natural History of Canada.
3The Province of Quebec Society for the Protection of Birds.
Quebec shows only three extant specimens from the
birds killed in Berthierville by Paquette in 1931: one of
unstated age and sex in a Berthierville school
museum, another of unstated age and sex in the
museum of College de Sainte-Anne-de-la-Pocatiére,
and an adult in the museum of the Deaf and Dumb
Institute (No. 31/16), “from a group of 4 or 5 which
had taken refuge in the church steeple.” This
annotation shows clearly that Gaboriault was con-
vinced that the species had not bred. None of the
extant specimens was a flightless young, and Paquette
never said that the birds had nested. Paquette may
have exaggerated the number of birds seen, and Créte
perhaps assumed that only breeding could explain
such a number of birds.
What appears to be the correct evaluation of the
record is found in Gaboriault’s notes. On a map of
Quebec he had marked records of occurrence with
different keyed symbols according to their seasonal
status. The symbol indicating the Berthierville record
on the map of the Barn Owl corresponds to a fall
visitor.
The records of occurrence to date (Table 1) show
only a pattern of casual vagrancy in the St. Lawrence
lowlands, with exceptional records eastward to
Kamouraska County (Mont-Carmel) and northward
in the Lake St. John area (Saint-Coeur-de-Marie).
The most recent record though suggestive, did not
provide conclusive evidence that the Barn Owls
actually bred within the limits of the province. A bird
in breeding condition was captured in a field near
Masson, Papineau County, only 1.5 km north across
the Ottawa River from the Ontario border; it was
seized by game wardens | or 2d later, on21 July 1977.
The specimen, now preserved in the National Mu-
seum of Natural Sciences of Canada (No. 67191), isa
female having a refeathered brood patch; its
reproductive tract contained 10 empty follicles and
several ova, the largest measuring 16 mm (Henri
Ouellet, personnal communication).
I thank Wilfrid Gaboriault who kindly lent me the
ornithological notes of his late brother. I also thank
Raymond Cayouette who allowed me to examine his
correspondence with F. Créte. Henri Ouellet reviewed
the manuscript. Victor Gaboriault died before he
could write a planned account on the distribution of
birds in Quebec; for this important contribution, he
should be considered as the author of this note.
Literature Cited
Cayouette, R. 1947. Barn Owl in Quebec. Auk 64: 631.
Godfrey, W.E. 1966. The birds of Canada. National
Museum of Canada Bulletin 203.
Paquette, A. 1961. J’ai gardé en captivité des oiseaux
rapaces. Jeune Naturaliste 11: 129-132.
Received 20 November 1978
Accepted 24 February 1979
1979
NOTES
325
Communal Roosting of Song Sparrows under Snowbank
MARTIN K. MCNICHOLL
Beak Consultants Ltd., 3530 11A St. N.E., Calgary, Alberta T2E 6M7
MeNicholl, Martin K. 1979. Communal roosting of Song Sparrows under snowbank. Canadian Field-Naturalist 93(3):
325-326.
Fifteen Song Sparrows ( Me/ospiza melodia) were found roosting under an ice-encrusted snowbank on Prince Edward Island.
The birds appeared to be keeping warm by a combination of roosting under snow and reducing individual distance.
Key Words: Individual distance, Prince Edward Island, Song Sparrow, sub-nivean environment, winter survival.
At 06:55 on 7 April 1977 I was about to cross a
stretch of snowbank 50 m long by | m wide in a ditch
in front of a farmhouse near French River, Prince
Edward Island, when 15 Song Sparrows ( Melospiza
melodia) emerged in rapid succession from a hole
approximately 4 cm in diameter in the bank. Exam-
ination of the hole showed that it led via a short tunnel
(approximately 10 cm long) to a small cavity, ap-
proximately 18-20 cm long by 11-14 cm wide, and
8 cm high, with no other exit. Previous melt had
caused the snowbank to become encrusted with ice.
The temperature was —2°C at the time, but with strong
north winds felt much colder. (I was uncomfortably
cold in an “Eskimo-style parka in which I am
normally warm in -20°C weather in the prairie
provinces.)
Nice (1943, p. 106) stated that Song Sparrows roost
in “weeds, hedges and small evergreens,” and I have
found no references to their roosting in holes or under
snowbanks. Sub-niveal roosting has been recorded
for Common Redpolls (Carduelis flammea) (Cade
1953: Novikov 1972), Dark-eyed Juncos (Junco
hyemalis) (Linsdale 1928), Tree Sparrows (Spizella
arborea) (Thompson 1934), Snow Buntings ( Plectro-
phenax nivalis) (Bagg 1943), and several Eurasian
passerines (Novikov 1972). In a recent summary of
several earlier accounts of passerines roosting under
the snow in Finland and the Soviet Union, Novikov
(1972) concluded that several species which regularly
winter in snow-covered areas frequently roost under
snow. Unlike most of the above-mentioned species,
Song Sparrows do not usually winter in snow-covered
areas, although many return to northern nesting areas
before all snow is gone, and there are many records of
individual birds wintering in northern areas.
Kendeigh (1961) demonstrated the energetic ad-
vantage to birds of roosting in cavities, and the well
known insulative properties of snow (Formozov 1946;
Elsner and Pruitt 1959) would enhance such an
advantage. This insulative ability of snow, however, is
poorest when the snow has been penetrated by water
or converted to ice, as in the present case. Thus, even
under the snowbank, the sparrows may have been
under considerable cold stress, a condition known to
induce breakdown of individual distance, resulting in
clumping together (Beal 1978). Although such crowd-
ing together is reported for several colonial or flocking
species in cold weather (e.g., Grubb 1973; Meservey
and Kraus 1976), | am not aware of reports of such
behavior in a species such as the Song Sparrow which
shows intraspecific intolerance all year (see Nice
1943). As territorial behavior tends to be particularly
strong shortly after return to the nesting area in
spring, this observation of communal roosting ap-
pears particularly remarkable.
Thus, the sparrows I observed appear to have
combined the strategies of roosting under snow and
clumping together to keep warm. In such a territorial
species, such a breakdown of individual distance
likely occurs only rarely.
I thank Anthony J. Erskine, William O. Pruitt, Jr.,
and two anonymous referees for helpful comments on
the manuscript.
Literature Cited
Bagg, A.M. 1943. Snow Buntings burrowing into snow-
drifts. Auk 60: 445.
Beal, K.G. 1978. Temperature-dependent reduction of
individual distance in captive House Sparrows. Auk 95:
195-196.
Cade, T. J. 1953. Sub-nival [sic] feeding of the Redpoll in
interior Alaska: a possible adaptation to the northern
winter. Condor 55: 43-44.
Elsner, R. W. and W. O. Pruitt, Jr. 1959. Some structural
and thermal characteristics of snow shelters. Arctic 12:
20-27.
Formozoy, A. N. 1946. Snow cover as an integral factor of
the environment and its importance in the ecology of
mammals and birds. Boreal Institute, University of
Alberta, Edmonton, Occasional Publication No. 1.
(Translated by W. Prychodko and W. O. Pruitt, Jr.)
Grubb, T. C., Jr. 1973. Absence of “individual distance”
in the Tree Swallow during adverse weather. Auk 90:
432-433.
326
Kendeigh, S. C. 1961. Energy of birds conserved by roost-
ing in cavities. Wilson Bulletin 73: 140-147.
Lindsdale, J. M. 1928. Birds of a limited area in eastern
Kansas. University of Kansas Science Bulletin 73: 517-
626.
Meservey, W.R. and G. F. Kraus. 1976. Absence of “in-
dividual distance” in three swallow species. Auk 93:
177-178.
Nice, M. M. 1943. Studies in the life history of the Song
Sparrow. Volume II. Transactions of the Linnaean
THE CANADIAN FIELD-NATURALIST
Vol. 93
Society of New York 6.
Novikov, G. A. 1972. The use of under-snow refuges
among small birds of the sparrow family. Aquilo Ser
Zoologica 13: 95-97.
Thompson, S. L. 1934. Unusual roosting of Tree Sparrow.
Canadian Field-Naturalist 48: 142-143.
Received 2 January 1979
Accepted 6 March 1979
Status of Eastern White Cedar, Thuja occidentalis, in Western
Nova Scotia
GORDON S. RINGIUS
Department of Botany, University of Alberta, Edmonton 16G 2E9
Ringius, Gordon S. 1979. Status of Eastern White Cedar, Thuja occidentalis in western Nova Scotia. Canadian Field-
Naturalist 93(3): 326-328.
The occurrence of Eastern White Cedar, Thuja occidentalis, in western Nova Scotia, in doubt since the initial report published
in 1877, is confirmed. A brief account of four of the five known stands in Cumberland County is presented and a map showing
the distribution of the species in Nova Scotia is included.
Key Words: Thuja occidentalis, rare, Cumberland County, distribution.
The earliest record of Eastern White Cedar, Thuja
occidentalis, from western (= Cumberland County)
Nova Scotia was published in 1877 by Lindsay.
Confirmation of Lindsay’s report has not been forth-
coming, however, and as a result, subsequent authors
(Table 1) have either cited Lindsay (Bentley and Smith
1962), included it without explanation (Maher et al.
1978: Saunders 1970), suggested that the species may
be present (Roland 1947), or have excluded it (Loucks
1962; Roland and Smith 1969). The purpose of this
note is to confirm the presence of Thuja occidentalis in
western Nova Scotia and to indicate where the known
stands are located.
On | April 1977, I observed a small stand of Thuja
occidentalis along the Trans-Canada highway about
5.8km E of the River Phillip - Oxford junction
(45°43’N, 63°50’W). The stand was located near the
base of a west-facing slope adjacent to a Speckled
Alder (Alnus rugosa) — Tamarack (Larix laricina)
swamp and consisted of less than 50 mature trees
scattered over an area of approximately 5 ha. The
trees were sparsely branched, many had single and
double bifurcate stems, and most showed reddish
brown discoloration of the lower and _ peripheral
foliage. The presence of numerous previous years’
cones indicated that the stand was producing seed.
Because snow still covered much of the ground,
however, no attempt was made to locate seedlings or
saplings. A voucher specimen was collected and has
been placed in the Smith Herbarium at Acadia
University (Ringius 796).
To determine whether other stands of Thuja occi-
dentalis have been located in western Nova Scotia, I
examined the collections at ACAD, DAL, DAO, and
NSPM (acronyms from Holmgren and Keuken 1974).
In addition, I contacted several people who are
involved with forestry in Nova Scotia. As a result of
these investigations, I discovered four additional
stands, and a previous unpublished account of the
stand described above. The locations of these stands
are shown in Figure |.
On file at NSPM are the following two reports
taken from the Chignecto Peninsula Inventory:
Eatonville near Apple River, Cumberland
Co., a cedar swamp less than one acre
[0.4 ha] in size. Sandy MacGregor,
Forestry Superintendent, Scott Paper
Company, New Glasgow.
A few miles southeast of Oxford, Cumber-
land Co., scattered cedar trees located
along the trans-Canada [highway] in this
area. Also a few cedar swamps of about 5
or 6 acres [2.0 — 2.4 ha] each between the
1979 NOTES 307
lec aa a5
47°
46° ‘\ —~
4) D S
Gig 6 ZA
4 » ire i a
\iror :
% .@ K \ ) ) 45
et
‘ ° Fp ean ~ X o
y Ss Thuja occidentalis
zh ‘
ie f LA @ Specimens Examined =|
44° Vv, f ‘ 44°
# © Other Reports
al SCALE
L ry 80 Kilometers
ge 64° 62° 60°
FIGURE 1. The distribution of Eastern White Cedar, Thuja occidentalis, in Nova Scotia, based on specimens examined at
ACAD, DAL, DAO, and NSPM, and on reports obtained through correspondence. A — Annapolis County,
C — Cumberland County, D — Digby County, K — Kings County, Y — Yarmouth County.
highway and the railroad. Cedar, which
used to be very common in Cumberland
Co., is now rare and there are only a few
cedar stands remaining. Department of
Lands and Forests, Oxford.
S. MacGregor (personal communication 1977)
stated that “the population of cedar at Apple River is
native as far as I know. There are only a few stunted
trees growing ona very tiny wet area.” Similarly, J. H.
Beattie, Supervisor Forest Resources, Department of
Lands and Forests, Oxford (personal communication
1977) stated the “cedar southeast of Oxford is native.”
He also said there are small stands at Lorneville,
Pugwash Junction, and near Joggins. All three sites
are in Cumberland County. The Lorneville stand was
at one time about 4 ha in size but has been seriously
reduced over the years. As a result of cutting in the
summer of 1977 only a few small trees of this stand
remain. On his suggestion 1 contacted Harold V.
Hatfield, Supervisor Forest Resources, District of
Cumberland West, Parrsboro, who was acquainted
with the Joggins stand. He provided information on
its location and referred to it as a “small amount of
wild cedar. [The trees] are young [and] scattered
among spruce and hardwood on an old burn.”
It is apparent that the stands of Thuja occidentalis
in Cumberland County are small, scattered, and rare.
In these respects they are similar to those in Anna-
polis, Digby, Kings, and Yarmouth counties in south-
western Nova Scotia where the species has been
known to occur since about 1801 (Table 1). In an
ecological study of the southwestern stands, Bentley |
and Smith (1962) found that although reproduction
was ample as indicated by seedling density, seedling
mortality was apparently high because few young
trees were present in any of the stands. They
concluded that competition from other tree species
was the limiting factor controlling the distribution of
Thuja occidentalis in Nova Scotia. It should be
mentioned that the species grows well enough when
planted; as an ornamental it is found throughout the
province with the possible exception of Cape Breton
Island.
I thank S. P. Vander Kloet of Acadia University,
Wolfville, Nova Scotia, for reviewing an early draft of
this note and offering helpful suggestions. Informa-
tion supplied by J. H. Beattie, H. V. Hatfield, and S.
MacGregor, and access to the various herbaria are
appreciated and gratefully acknowledged.
328 THE CANADIAN FIELD-NATURALIST Vol. 93
TABLE 1—Previous reports of Eastern White Cedar, Thuja occidentalis, in Nova Scotia
Author Date Locality
Smith! ca. 1801 Annapolis Valley (not plentiful)
Lindsay 1877 Cumberland County
Macoun 1886 Swamp within 3 mi[4.8 km] of Annapolis[ Annapolis County], but said to grow on
the Bay of Fundy near [Annapolis]; rare in Nova Scotia
Fernald 192] Digby County, Yarmouth County (lakesides)
Roland 1947 Annapolis Valley, Digby, Yarmouth counties; possible existence on the isthmus
between Nova Scotia and New Brunswick [Cumberland County]
Bentley and Smith 1962 Annapolis, Digby, Kings counties; reported from Cumberland County [cite
Lindsay 1877]
Loucks 1962 Wentworth Lake District [= portions of Digby and Yarmouth counties]; local
Roland and Smith 1969 Annapolis Valley [includes Kings County], Annapolis, Digby counties
Saunders 1970 Annapolis Valley, Digby, Yarmouth counties; slight ‘spillover’ from New
Brunswick on the Chignecto Isthmus [Cumberland County]
Maher et al. 1978 Annapolis, Cumberland, Digby, Kings, Yarmouth counties
'Cited by Gorham (1955).
Literature Cited
Bentley, P. A. and E. C. Smith. 1962. A study of northern
white cedar and jack pine in Nova Scotia. Proceedings of
the Nova Scotian Institute of Science 24: 376-398.
Fernald, M. L. 1921. The Gray Herbarium expedition to
Nova Scotia. Rhodora 23: 89-111.
Gorham, E. 1955. Titus Smith, a pioneer of plant ecology in
North America. Ecology 36: 116-123.
Holmgren, P. A. and W. Keuken. 1974. Index herbar-
iorum. Part I. The herbaria of the world. 6th edition.
Oosthoek, Scheltema, and Holkema, Utrecht, Nether-
lands.
Lindsay, A. W. H. 1877. A catalogue of the flora of Nova
Scotia. Proceedings of the Nova Scotian Institute of
Science 4: 184-222.
Loucks, O. L. 1962. A forest classification for the maritime
provinces. Proceedings of the Nova Scotian Institute of
Science 25: 85-167.
Macoun, J. 1886. Catalogue of Canadian plants. Part III.
Apetalae. Dawson Brothers, Montreal.
Maher, R. V., D. J. White, G. W. Argus, and P. A. Keddy.
1978. The rare vascular plants of Nova Scotia. Syllogeus
Series Number 18. 37 pp.
Roland, A. E. 1947. The flora of Nova Scotia. Proceed-
ings of the Nova Scotian Institute of Science 21: 97-644.
Roland, A. E. and E. C. Smith. 1969. The flora of Nova
Scotia. Nova Scotia Museum, Halifax. 743 pp.
Saunders, G. L. 1970. Trees of Nova Scotia. Department of
Lands and Forests, Nova Scotia. 102 pp.
Received 21 November 1978
Accepted 7 April 1979
1979
NOTES
329
Snowy Egret in the Northwest Territories
RICHARD W. QUINLAN
Canadian Wildlife Service, #1000, 9942 - 108 Street, Edmonton, Alberta T5K 2J5
Quinlan, Richard W. 1979. Snowy Egret in the Northwest Territories. Canadian Field-Naturalist 93(3): 329.
Key Words: Snowy Egret, Northwest Territories, first record.
On 23 June 1977, a Snowy Egret ( Egretta thula) was
seen near Fort Simpson (61°53’N, 121°25’W). This is
the first recorded sighting in the Northwest Territories
and the most northerly in North America. This species
breeds from the central United States through the
West Indies and Central America to Chile and
Argentina. It is a rare non-breeding wanderer to
Alberta, Saskatchewan, and British Columbia (W. E.
Godfrey, 1966. The birds of Canada. National
Museum of Canada Bulletin 203). There are six
records of Snowy Egrets in Alberta, all in May and
June (C.S. Houston and M.I. Houston, 1976.
American Birds 29: 74-77; W. R. Salt and J. R. Salt,
1976. The birds of Alberta. Hurtig, Edmonton); the
most northerly was at Sandy Lake (52°47’N, 111°00’W)
in May 1909. The most northerly of four Saskatche-
wan sightings was at Saskatoon (52°07’N, 106°38’W)
on 15 May 1977(E. M. Serr, 1977. American Birds 31:
1013-1016). Near Juneau, Alaska (58°20’N,
134°20’W), a Snowy Egret was observed 18-24 May
1957 (I. N. Gabrielson and F. C. Lincoln, 1959. The
birds of Alaska. Stackpole Co., Harrisburg, Pa.)
The bird reported here was observed on the
downstream (northwest) end of Fort Simpson Island
in the Mackenzie River. It was first sighted on 23 June
by Richard Quinlan and Yvonne Desilets, who
watched it for 45 min from a distance of approxi-
mately 50 m. Quinlan, with James Steele and (once)
Bruce Reilly, also saw the egret on each of the next
4d. All observations except the last (1 km down-
stream on a small island) were at the original location.
A super-8 movie film taken on 25 June was viewed
by Henri Ouellet of the National Museum of Natural
Sciences, who verified the identification. Bernard
Gollop provided a number of additional records from
Alberta and Saskatchewan.
Received 11 December 1978
Accepted 19 March 1979
Letters
Is Biology Unknown?
The recent Guest Editorial by Yorke Edwards,
“Biology — The Unknown Science?” (Canadian
Field-Naturalist 93(1): 6-9; 1979), invites a reply and
several counter-proposals. Of all the sciences, Biology
is probably the best known today. It has its Suzuki,
and the media are full of “gee-whiz” articles on genetic
engineering, tissue culture, test-tube babies, and
Cinderella crops. What is missing, according to
Edwards, is the synthesizing scientist who will put all
these marvelous bits and pieces together “so that
people may understand their world.”
There are a couple of serious problems with this
diagnosis. Implicit is the assumption that science is
engaged in discovering reality, not making it. But
suppose that at least some science (and certainly all
the technology that science nurtures) is motivated not
so much by a desire to understand as to control and
change? Then people may rightly question whether
scientists are all admirable, and whether every old
kind of science is A-OK. They might, for example, ask
what exactly biologists are up to when they study the
deep-diving abilities of seals, or the cold resistance of
wolves’ feet. “Why,” your innocent scientist will
answer, “we mean to help our fellow men to explore
safely the ocean floor for mineral treasures, and
warmly to run around on the snow while searching for
good Canadian oil!” “But surely,’ the layman
responds, “you investigate these things with an enrich-
ing sense of wonder and awe, and this you will
transmit to me so that I too may understand and
marvel?” “Will you please go away and don’t bother
me,” comes the reply, “I’m preparing a scientific
article (on which my next promotion and merit
increase depend) for one of the NRC journals and
tomorrow I’m off to Ottawa as a consultant to the
Department of National Defence.”
A related questionable assumption is that the
synthesis of knowledge needed to understand the
world will come from a change in attitude of those
who know the facts (the analytical scientists) rather
than from a newly constituted science that sets as its
goal the sympathetic understanding of world eco-
systems. After all, how realistic is it to expect nuclear
physicists to go to bat for Ecological Reserves, or
mining geologists to take up the cudgels for a
decentralized solar society? Their training is wrong,
and so is the training of most biologists. It is not that
Biology is unknown but that the proper subject is
unknown.
Finally, we hear much today about morality in
science, meaning that no matter what is buzzing
around in their Pandora’s box, scientists should
carefully explain to the public the undoubted advan-
tages of taking off the lid. Such morality is largely self-
serving. There needs to be, as Aldo Leopold said long
ago, a land ethic, acommitment to something external
to the human race, something equally if not more
important. The main problem with science, and the
reason for the lack of Earthcare, is the narcissism of
the human race.
J. STAN ROWE
9 March 1977
Department of Plant Ecology, University of Saskatchewan,
Saskatoon, Saskatchewan 2N4 0WO
Never one to be uninterested in reactions to what I
write, I gladly accept this opportunity to comment on
Dr. Rowe’s thoughtful comments, and I do so briefly.
] read them with complete sympathy and agreement,
for most of his thoughts are not far from those I hold,
or have held.
Inevitably the world we see is the part of it that we
are in, and my current views must therefore be colored
by a long association with government and primary
industry. I can only say in addition to what I have said,
that I find my part of the world quite frightening
because of the values used in deciding destructions of
the living parts of Earth as well as of the mineral bases
for that life. As Dr. Rowe probably knows, my copy of
Leopold is as old and as well thumbed as his. In sucha
brotherhood as this, there can be only relatively minor
differences in views of land and its life.
YORKE EDWARDS
30 March 1979
British Columbia Provincial Museum, Victoria, British
Columbia V8V 1X4
330
Tribute to HOYES LLOYD, 1888-1978
Hoyes Lloyd, who died in his 89th year in Ottawa
on 21 January 1978, was the central figure in
Canadian nature conservation activities during the
critical 25 years from 1918 to 1943. During that
period, nature conservation grew from a personal
interest in the minds of a few people toa responsibility
taken seriously by government. The first person given
a professional role in nature conservation by the
Canadian Government, Lloyd carefully expanded
and consolidated activities aimed at the conservation
of migratory birds and of wildlife in the national parks
and the Territories, laying the groundwork for the
establishment of the Canadian Wildlife Service, which
was formed (as the Dominion Wildlife Service) in
1947. His accomplishments rested upon his thorough
knowledge of wildlife, his keen sense of what was
possible and practical and, perhaps most importantly,
his ability to pick a path through political and
bureaucratic tangles with calmness, diligence, and
humor.
For anyone of Hoyes Lloyd’s generation, there was
scant opportunity for a professional career in wildlife
conservation. Almost all of those who broke into the
field before 1930 were first obliged to seek means of
livelihood less closely matched to their real interests.
Lloyd, born in Hamilton in 1888, and educated at
Harbord Collegiate Institute in Toronto and at the
University of Toronto, found his first full-time
employment in applied chemistry. From 1909 to 1911,
while working for his Master’s degree in Chemistry, he
served as an assistant in the department, and during
1911 and 1912 he was employed by a manufacturer of
chemicals. He entered public service in 1912 when he
was appointed chemist in charge of milk control for
the City of Toronto, and a few years later he played an
important role in bringing about compulsory pas-
teurization of the city’s milk supply. Not until 1918
was he able to give his full time to conservation.
But Lloyd was an active student of natural history
long before that. Just after the turn of the century, he
began to keep records of the occurrence and behavior
of birds and to collect plants and birds. During the
next few years he roamed the woods, fields, and
marshes around Toronto, often with young friends
who shared his interests, building up his knowledge of
nature and through his collections establishing per-
manent records of the occurrence of birds and plants.
In 1903 he happened ona copy of The Auk ina book
store and learned that bird study was a serious thing
pursued by serious persons. In 1904, 1905, and 1906 he
won prizes for plant collections entered in the
Canadian National Exhibition. In the summer of
1909, which he later described as one of the most
exciting times of his life, he hiked and paddled the
wilderness as a forest ranger and deputy game warden
on the Temagami Forest Reserve. In the same year he
won a gold medal at the Canadian National Exhibi-
tion for his collection of bird skins.
Full-time employment as a chemist in the field of
public health from 1917 to 1918 did not diminish his
activities as a field naturalist. During that period he
continued to observe and collect. In 1916 he became
an Associate Member of the American Ornithologists’
Union and in 1917 he wrote “Ontario Bird Notes” his
first paper for The Auk.
Lloyd’s opportunity to work in the field he loved
came as attitudes to nature and the use of natural
resources began to change. Early in the century, a few
North Americans began to see the folly of profligate
use of resources. In Canada, the Commission on
Conservation issued its series of thoughtful reports in
the years 1910-1913. Among its recommendations
was one that called for an agreement between Canada
and the United States for the protection of migratory
birds. Discussions and negotiations proceeded, lead-
ing to the signature of the Migratory Birds Con-
vention in 1917, and the enactment of enabling
legislation in Canada, the Migratory Birds Conven-
tion Act in 1918. Within the year Hoyes Lloyd wona
competition to head the one-man Migratory Birds
Unit in the Department of the Interior.
Lloyd remained with the Department of the
Interior and its successor, the Department of Mines
and Resources, for 25 years, in charge of a wildlife unit
that was gradually given broader responsibilities but
was always too small to meet its responsibilities to his
satisfaction. The early years were particularly diffi-
cult: the task immense, the resources infinitesimal. At
first Lloyd served as policy-maker, field man, chief —
clerk, and everything in between, but within three
years he had a clerical assistant and three migratory
bird officers to help him cover all of Canada. An
urgent requirement then as now was to tell people
about the Migratory Birds Convention Act, and the
North West Game Act, for the administration of
which he was also responsible, and to persuade them
that the Acts and Regulations must be observed. This
meant writing letters and leaflets; meeting sportsmen
and naturalists formally and informally; advising
teachers, wardens, police and magistrates; and con-
ferring with officials of other government depart-
ments, provincial governments and the Government
of the United States. It also meant travelling from one
end of Canada to the other to familiarize himself with
conservation problems in the field, of which there
were many, and frequent meetings with his superiors
331
332
in Ottawa to seek more support for the work that had
to be done. Progress was slow but steady. Migratory
bird sanctuaries were established; the Migratory Bird
Regulations were improved and their enforcement
became more effective; cooperation with the United
States in studying the migrations and populations of
migratory birds was initiated and developed.
In 1922, Hoyes Lloyd organized and served as
secretary to the first Federal-Provincial Wildlife
Conference, an institution that soon came to meet
annually and enabled effective exchange of informa-
tion and views among the wildlife authorities of all the
provinces and the federal government. The “Federal-
Provincials,” which are still held each year, have been
highly effective in the development of wildlife con-
servation in Canada. At that first conference Lloyd no
doubt displayed the capacity for unobtrusive but
effective group leadership that he was later to employ
so often in so many different settings. He was
President of The Ottawa Field-Naturalists’ Club from
1923 to 1925; President of the International Associa-
tion of Game, Fish and Conservation Commissioners
in 1929-30; Chairman of the Canadian Section of the
International Council for Bird Preservation from
1927 to 1954 and Vice-President of the Council itself
from 1938 to 1950. He regularly attended and, as the
senior Canadian official, was often a key-note speaker
at the annual meetings first known as the North
American Game Conference (he was chairman in
1935) and now as the North American Wildlife and
Natural Resources Conferences. He organized and
chaired innumerable less formal meetings in many
parts of Canada and in the United States.
By 1943 when Lloyd retired after 25 years of public
service, wildlife protection in Canada was widely if
not universally accepted and wildlife management asa
more broadly based and purposeful discipline was
beginning to emerge. Much of the credit for that
development is his.
Retirement gave Hoyes Lloyd the opportunity to
extend his professional interest in birds. This had not
been neglected in his years in the Civil Service; he was
increasingly active in the American Ornithologists’
Union during the 1920s and 1930s and his bird
collection slowly continued to grow. In 1942 he
became a Vice-President of the Unionand he served as
its President from 1945 to 1948. His participation in
the work of the International Council for Bird
Preservation became more active in his post-retire-
ment years during which he attended and contributed
actively to each of its quadrennial conferences until
1972. All this work was, of course, voluntary; though
he often represented the Canadian Government
abroad in places suchas Helsinki, Caracas, Bulawayo,
and Tokyo, he paid his own way and he did so without
complaint. Old associates in the American Ornitholo-
THE CANADIAN FIELD-NATURALIST
Vol. 93
gists’ Union and the Wildlife Society continued to
welcome him to their meetings which he attended until
1973.
Even after travel became less easy for him Lloyd
kept up an active correspondence with his many
friends and received occasional contributions to his
collections. His collections of plants, birds, and
mammals along with his field journals covering a
period of 71 years and his extensive library, are now in
the care of the Royal Ontario Museum.
Hoyes Lloyd wrote over 120 papers and notes for
publication. Of these, over 100 were contributions to
The Canadian Field-Naturalist; he did much to
develop its reputation as a worthy source of informa-
tion about Canadian wildlife. In 1944 he published in
our journal a 32-page annotated list “The Birds of
Ottawa,” which consolidated and added to notes on
Ottawa birds published in 1923, 1924, 1925, 1932,
1936, and 1937. In 1945, 1949, and 1951 he again
added to his regional list. This work provided a major
reference for the bird group of The Ottawa Field-
Naturalists’ Club and contributed occurrence records
to many later works on the distribution of North
American birds. Lloyd’s bibliography includes papers
dealing with many other records of bird distribution
in other parts of Canada as well as records of
migrations derived from the recoveries of banded
birds and accounts of the introduction of Ring-necked
Pheasant and Hungarian Partridge to various parts of
Canada.
The only Canadian to be the recipient of the two
premier honors in the wildlife field in North America
was Hoyes Lloyd. He was presented with the Leopold
Award by the Wildlife Society in 1956 and the Seth
Gordon Award by the International Association of
Fish, Game and Conservation Commissioners in
1974. He was made an Honorary Member of both
those organizations and he was elected a fellow of the
American Ornithologists’ Union and of the New York
Zoological Society. He was an Honorary life member
of the Quebec Zoological Society, the Outdoor
Writers Association of America, and The Ottawa
Field-Naturalists’ Club.
Lloyd was married in 1913 to Wilmot Lockwood of
Memphis, Tennessee. His two daughters and his son,
and later his grandchildren and great-grandchildren
all lived in Ottawa, and with his wife and himself
formed a close-knit group that was rarely long
separated. Their home in Rockcliffe Park, selected
because he heard a Wood Thrush sing when he first
inspected the spacious grounds, provided an ideal site
for frequent family parties.
A warm and friendly person, unfailingly courteous,
Hoyes Lloyd was always generous with his time and
knowledge, particularly with younger people. It was
often my personal pleasure to benefit from his counsel
HOyYES LLOYD, 1888-1978 333
1979
TTT ATEN
jp L/h Hi
WOT
Hoyes Lloyd
On the occasion of his receipt of the 1974 Seth Gordon Award, the highest honor given by the International Association of
Game, Fish and Conservation Commissioners, for his outstanding work in wildlife conservation.
334
THE CANADIAN FIELD-NATURALIST
and encouragement. While he knew that the continu-
ing progress of nature conservation would be well
served by the help he gave to those who would take up
his cause, his personal relationships went far beyond
such mere calculation. He truly loved his fellow man.
DAVID A. MUNRO
IUCN (International Union for Conservation of Nature and
Natural Resources), 1110 Morges, Switzerland.
Publications of Hoyes Lloyd
1917
1918
1918
1919
1919
1919
192]
1921
1921
192]
1922
1922
1922
1922
1922
1922
1923
1923
1923
1923
1923
1923
1924
1924
1924
Ontario bird notes. Auk 34: 457-460.
Spring in Birdland. Youth and Services. C.S.E.T.
Department of Ontario, Volume 20(32).
The extraction of fat from bird-skins.
164-169.
The Ornithological collector and the law. Canadian
Field-Naturalist 33: 93-94.
Permits to collect birds for scientific purposes in
Canada. Auk 36: 621-623.
Protection of bird neighbours.
Leaflet. March.
Bird notes by the way in the Maritime Provinces.
Canadian Field-Naturalist 35: 11-13.
The Town of Yarmouth, N.S. buys a bird sanctuary.
Canadian Field-Naturalist 35: 35-36.
An aquatic habit of the pigeons. Canadian Field-
Naturalist 35: 98-99.
Attracting birds to the prairie home. Canadian
Forestry Magazine 17: 76-78.
Some of Captain Henry Toke Munn’s observations on
birds of Baffin Island and vicinity. Canadian Field-
Naturalist 36: 49-50.
Larvae of Phormia chrysorrhea Macq. found upon
nestling Bluebirds. Canadian Field-Naturalist 36:
116.
Review of “Some useful Australian birds.” Canadian
Field-Naturalist 36: 118-119.
A recent breeding record of the Trumpeter Swan in
Alberta. Canadian Field-Naturalist 36: 143-144.
Popular names for birds. Auk 39: 144.
Some northern records of the Turkey Vulture.
Canadian Field-Naturalist 36: 178-179.
Protecting nesting sites. Auk 40: 571.
Observations on the wintering flocks of Canada Geese
in Nova Scotia. Canadian Field-Naturalist 37: 26-28.
Brown Creeper the victim of strange accident.
Canadian Field-Naturalist 37: 75.
The birds of Ottawa. Canadian Field-Naturalist 37:
101-105; 125-127; 151-156.
The eating of birds by chipmunks. Canadian Field-
Naturalist 37: 118.
Loss of Whistling Swans at Niagara Falls. Canadian
Field-Naturalist 37: 138.
The birds of Ottawa. Revised to March 20, 1923.
Canadian Field-Naturalist 38: 10-16.
Christmas bird census 1923. Canadian
Naturalist 38: 17.
An unusual sight description of a bird verified.
Canadian Field-Naturalist 38: 38.
Auk 35:
Dominion Parks
Field-
1924
1924
1924
1924
1924
1925
1925
1925
1925
1925
1925
1925
1925
1925
1925
1926
1926
1926
1926
1926
1926
1926
1927
1927
1927
1927
1927
Vol. 93
A flock of Trumpeter Swans. Canadian Field-
Naturalist 38: 57-58.
Another mystery bird. Canadian Field-Naturalist 38:
76.
Field Sparrow and Towhee at Christie Lake, Ontario.
Canadian Field-Naturalist 38: 180.
An old breeding record for the Herring Gull at Lac des
Iles, Labelle Co., Quebec. Canadian Field-Naturalist
38: 193-194.
A Fulmar at Arnprior, Ontario. Canadian Field-
Naturalist 38: 194.
The birds of Ottawa — Addenda to October 2, 1924.
Canadian Field-Naturalist 39: 16-17.
Mystery bands. Canadian Field-Naturalist 39: 43-44.
The recording of acclimatization experiments.
Canadian Field-Naturalist 39: 63-64.
A second record of Scissor-tailed Flycatcher at York
Factory, Manitoba. Canadian Field-Naturalist 39:
86.
A juvenile Richardson’s Owl from Grand Manan, New
Brunswick. Canadian Field-Naturalist 39: 113.
A mystery band solved. Canadian Field-Naturalist
39: 114.
The acclimatization of the Fox Squirrel at Pelee
Island, Ontario. Canadian Field-Naturalist. 39: 138.
The introduction of the Muskrat and the Rocky
Mountain Goat on Vancouver Island. Canadian
Field-Naturalist 39: 151-152.
The introduction of the European Grey Partridge in
Manitoba. Canadian Field-Naturalist. 39: 172.
Christmas bird censuses in Canada. Canadian Field-
Naturalist 39: 173.
Christmas bird census
Naturalist 40: 12-13.
Yellow-bellied Sapsucker infested with tape worms.
Canadian Field-Naturalist 40: 67.
More mystery birds. Canadian Field-Naturalist 40:
88.
Canadian Government motion picture films showing
natural history subjects. Canadian Field-Naturalist
40: 100-102.
Review of “Animal life in the Yosemite” by Joseph
Grinnell and Tracy Irwin Storer. Canadian Field-
Naturalist 40: 114-115.
The 44th stated meeting of the American Orni-
thologists’ Union, Ottawa, October 11-17, 1926.
Canadian Field-Naturalist 40: 189-190.
Review of “The relation of birds to woodlots in New
York State’? — W.L. McAtee. Canadian Field-
Naturalist 40: 194.
A mystery solved. Bulletin of the Northeastern Bird-
Banding Association 3: 17-18.
Point Pelee should have a bird banding station.
Canadian Field-Naturalist 41: 89-90.
Identification of flies found infesting Bluebirds.
Canadian Field-Naturalist 41: 111.
Transfers of Elk for re-stocking. Canadian Field-
Naturalist 41: 126-127.
Additional comments regarding Mallards wintering at
Buffalo Lake, Alberta, 1923-24. Canadian Field-
Naturalist 41: 138.
1925. Canadian Field-
IST)
1927
1927
1927
1927
1927
1928
1928
1928
1928
1928
1929
1929
1929
1930
1930
1931]
1932
1932
1933
1933
1933
1933
1934
1934
1935
1936
1936
1937
HoyYES LLOYD, 1888-1978
Another mystery band. Canadian Field-Naturalist 41:
139.
Rocky Mountain Sheep transferred from Banff to
Spence’s Bridge, B.C. Canadian Field-Naturalist 41:
140.
A mystery band. Canadian Field-Naturalist
155-156.
Work of the Canadian Section, The International
Committee for Bird Protection. Canadian Field-
Naturalist 41: 156.
European Widgeon crosses from Iceland. Canadian
Field-Naturalist 41: 171.
Notes on the introduction of the pheasant in some of
the provinces of Canada. Canadian Field-Naturalist
42: 143-145.
The spread of the European Starling. Canadian Field-
Naturalist 42: 178.
Bird banding in Japan. Canadian Field-Naturalist 42:
178.
Trans-Atlantic banded pigeon recovery. Canadian
Field-Naturalist 42: 178.
A method for cleaning large bird skins. Canadian
Field-Naturalist 42: 207-208.
Review of “Birds of Western Canada.” 2nd Edition. By
P. A. Taverner. Canadian Field-Naturalist 43: 109.
What surgeon to the birds treated this Pine Grosbeak?
Canadian Field-Naturalist 43: 167-168.
Who banded this Gull? Canadian Field-Naturalist 43:
189.
Whistling Swan at Rigaud, Quebec. Canadian Field-
Naturalist 44: 167.
Henderson’s “The practical value of birds.” Auk 47:
302-304.
Progress of conservation in Canada. Bird Banding II:
105-115.
Starlings bathing in
Naturalist 46: 22.
The birds of Ottawa — Addenda. Canadian Field-
Naturalist 46: 123-127, 162-166.
An occurrence of the Clay-coloured Sparrow, Spizella
pallida, in Renfrew County, Ontario. Canadian Field-
Naturalist 47: 36.
Wild Geese killed by lightning. Canadian Field-
Naturalist 47: 72-73.
Obituary Norman Criddie
Field-Naturalist 47: 145-147.
Review of “Game management” by Aldo Leopold.
Canadian Field-Naturalist 47: 159-160.
The conservation of wild bird life. Beaver, Outfit 265,
June 1934. p. 22.
An Ontario sand-fall. Canadian Field-Naturalist 48:
93-95.
Review of “Sheep and bear trails” by John P. Holman.
Canadian Field-Naturalist 49: 152.
The late Norman Criddle’s record of the Snowshoe
Rabbit (Lepus americanus) at Aweme, Manitoba.
Canadian Field-Naturalist 50: 129-130.
The birds of Ottawa — Addenda to February 28, 1935.
Canadian Field-Naturalist 50: 143-144.
A nest-hunting Broad-winged Hawk. Canadian Field-
Naturalist 51: 12.
41:
winter. Canadian Field-
1875-1933. Canadian
1937
1937
1937
1937
1938
1939
1939
1939
1939
1939
194]
1943
1943
1943
1944
1944
1945
1948
1949
1951
1951]
1951
1953
1953
1953
1954
1955
1956
335
Short-eared Owl (Asio flammeus flammeus) at Baker
Lake, N.W.T. Canadian Field-Naturalist 51: 13.
A wintering Towhee at Ottawa. Canadian Field-
Naturalist 51: 77.
Banded European Widgeon recovered in Prince
Edward Island. Canadian Field-Naturalist 51: 77.
Twenty year old Ferruginous Rough-legged Hawk.
Canadian Field-Naturalist 51: 137.
Some aspects of Canadian Bird-life. Canadian Geo-
graphic Journal 16: 265-283.
Review of “Canoe country” by Florence Page Jacques.
Canadian Field-Naturalist 53: 13.
Banded Kittiwake from Russia recovered in New-
foundland. Canadian Field-Naturalist 53: 29-30.
Andrew Halkett, naturalist, 1854-1937. Canadian
Field-Naturalist 53: 31-32.
Lorene Squire studies prairie birds. Canadian Geo-
graphic Journal 18: 19-25.
Deciphering numbers on much-worn bird bands.
Canadian Field-Naturalist 53: 131.
A Kittiwake Gull (Rissa tridactyla) banded near
Murmansk, U.S.S.R., recovered in Newfoundland.
Canadian Field-Naturalist 55: 138.
Review of “Ornithologists of the United States Army
Medical Corps: Thirty-six biographies” by Edgar
Erskine Hume, Colonel. Canadian Field-Naturalist
57: 14.
Review of “Wildlife of Idaho” by William M. Rush.
Canadian Field-Naturalist 57: 14-15.
Dr. T. Gilbert Pearson. Canadian Field-Naturalist 57:
141.
Review of “Some aspects of Canadian birds” by A. L.
Rand. Canadian Field-Naturalist 58: 18.
The birds of Ottawa. Canadian Field-Naturalist 58:
143-175.
Birds of Ottawa, 1944, an inadvertent omission.
Canadian Field-Naturalist 59: 150.
Review of “Wings in the wilderness” by Allan D.
Cruickshank. Canadian Field-Naturalist 62: 166.
Bird records for the Ottawa District. Canadian Field-
Naturalist 63: 31-34.
Duck Hawk at Blue Sea Lake, Quebec. Canadian .
Field-Naturalist 65: 46.
James Bay Sparrow at Ottawa. Canadian Field-
Naturalist 65: 46.
Recent Ottawa District bird records. Canadian Field-
Naturalist 65: 46.
Dead Golden Eagle at Perkins
Canadian Field-Naturalist 67: 44.
Review of “Proceedings of the Xth International
Ornithological Congress, Uppsala, June 1950.”
Canadian Field-Naturalist 67: 45-46.
Wilson’s Petrel, Oceanites oceanicus oceanicus (Kuhl)
at Lake Deschenes, Quebec. Canadian Field-
Naturalist 67: 140.
Clyde Louis Patch, 1887-1952. Canadian Field-
Naturalist 68: 125-126.
Mid-Atlantic migration of Long-tailed Jaegers and
Terns. Canadian Field-Naturalist 69: 24.
Ralph Emerson DeLury, 1881-1956,
Canadian Field-Naturalist 70: 169-170.
Mills, Quebec.
obituary.
336
1959
1961
1961
1963
1963
1965
1920
1921
THE CANADIAN FIELD-NATURALIST
William Henry Lanceley (1893-1958). Canadian
Field-Naturalist 73: 183.
A Manitoba occurrence of the Black-headed Gros-
beak. Canadian Field-Naturalist 75: 109.
Faith Fyles, artist-naturalist, 1875-1961. Canadian
Field-Naturalist 75: 220.
Review of “The cry of a bird” by Dorothy Yglesias.
Canadian Field-Naturalist 77: 124.
Catharacta skua Brunnich sighted in North Pacific.
Canadian Field-Naturalist 77: 127.
A review of Thoreau on birds.
Naturalist 79: 261.
J. H. Fleming and Hoyes Lloyd. Ontario bird notes.
Auk 37: 429-439.
Hoyes Lloyd and F.C. Hennessey. Annual 1920
Christmas bird census for Ottawa and Hull. Bird Lore
7p Gy).
Canadian Field-
1929
1948
1950
Vol. 93
Hoyes Lloyd and Norman H. H. Lett. Obituary —
George Rivers White, 1856-1927. Canadian Field-
Naturalist 43: 103-104.
Hoyes Lloyd and Robert G. Lanning. Observations on
the birds of Renfrew County, Ontario. Canadian
Field-Naturalist 62: 47-65.
Hoyes Lloyd, Robert G. Lanning, and Sheldon Davis.
Additional notes on the birds of Renfrew County,
Ontario. Canadian Field-Naturalist 64: 140-142.
Compiled by V. M. HUMPHREYS
National Museum of Natural Sciences, National Museums
of Canada, Ottawa, Ontario
KIA 0M8.
News and Comment
Robert Lister Receives Award
The fourth Loran L. Goulden Memorial Award for
outstanding work on natural history within the
province of Alberta was awarded to Robert Lister at
the annual meeting of the Edmonton Natural History
Club on 9 January 1979. Bob Lister, Honourary
President of the Edmonton Bird Club, served as an
indispensible assistant to the late Prof. William
Rowan for many years, while the latter conducted his
world-renowned experiments on influence of photo-
period on migration; however, he is recognized in this
award primarily for his long-term role in encouraging
other naturalists in Alberta, through his newspaper
column, his role in founding the Edmonton Bird Club,
his social activities, and other means. His forthcoming
book on the birds of Beaverhill Lake, based largely on
Rowan’s work, marks a fitting occasion on which to
recognize Bob through this award. Further details on
his career and a list of his publications will appear ina
future issue of the Alberta Naturalist.
Martin K. McNicholl
Chairman, Loran L. Goulden Memorial
Award Selection Committee
Notice of Change to the By-laws of The Ottawa Field-Naturalists’ Club
A motion to change By-law 15 of The Ottawa Field-Naturalists’ Club was passed unanimously by the Council at
the meeting of 12 February 1979. This By-law now reads as follows:
15. Membership Dues
The schedule of dues shall be as follows:
Individual $10.00
Family $12.00
Sustaining $25.00
Life Membership shall be granted upon payment of a single sum of $200.00.
Article or Note?
For The Canadian Field- Naturalist the assigning of
an accepted manuscript to one category or the other is
currently done solely on the basis of the manuscript’s
length. We do not differentiate Articles from Notes on
the basis of the quality or importance of the
observations or research, although these criteria are
stated or implied for some other scientific journals.
Notes are run on one after the other so as not to be
wasteful of available costly space because we are
charged on a per page basis for every page that
contains any printing whatsoever. Sometimes referees
and editors request that authors condense their
papers. The aim of the constructive advice is that the
new information is presented as clearly and concisely
as possible. Thus if an author focuses on the objectives
of his or her study and the purpose of his or her report,
Diana R. Laubitz, Recording Secretary
the manuscript will be more succinct and its message
more easily grasped.
Beginning with the 1977 issues, I eliminated the
main difference between the earlier printed versions of
Articles and Notes by starting to use the same basic
format for both. With this issue I have further
eliminated the previous difference in type sizes.
Therefore, shorter papers will be published as Notes
and longer ones as Articles but they will both be in the
same format. No differentiation will be made by the
Editor nor should be construed by authors or readers
regarding the relative significance of the Notes as
compared to Articles in The Canadian Field-Natur-
alist.
LORRAINE C. SMITH,
Editor
337)
338 THE CANADIAN FIELD-NATURALIST
Voly93
Notice of Motion to Amend the Constitution of The Ottawa Field-Naturalists’ Club
Notice of a motion to amend the Constitution of
The Ottawa Field-Naturalists’ Club was received, in
accordance with Article 23 of the Constitution, for
presentation at the next Annual Business Meeting. It
is proposed that: Article 11, now reading:
“AUDITORS. Two Auditors shall be elected
by open vote at the Annual Business Meeting.
They shall examine the Treasurer’s accounts
and certify as to their correctness.”
be amended to read as follows:
AUDITOR. An Auditor shall be elected by
open vote at the Annual Business Meeting. The
Auditor shall examine the Treasurer’s accounts
and certify as to their correctness.
Motion proposed by Courtney Gilliatt, seconded by
William Cody. This motion is in accordance with a
recommendation made at the last annual meeting.
Diana R. Laubitz,
Recording Secretary
Request for Participants — International Shorebird Surveys 1979
A cooperative International Shorebird Survey
scheme was organized by the Canadian Wildlife
Service and the Manomet Bird Observatory in 1974 to
obtain information on shorebird migration and to
identify and document areas of major importance.
Much very valuable information has come from
contributors throughout eastern Canada and the
USA, the Caribbean Islands, and Central and South
America, and this is being used in assessing require-
ments for the future protection and conservation of
the birds and their habitat. Continuation of the project
is planned for 1979. Any observer who will participate
in regular counts of shorebirds during migration
periods, as well as during the winter in shorebird
wintering areas, 1s asked to contact one of the
undersigned. Occasional counts from observers visit-
ing shorebird areas on an irregular basis would also be
most welcome. For areas in Canada: Dr. R. I. G.
Morrison, Canadian Wildlife Service, 1725 Wooa-
ward Drive, Ottawa, Ontario, Canada KIG3Z7. For
areas in USA, Caribbean Islands, Central and South
America: Brian A. Harrington, Manomet Bird Ob-
servatory, Manomet, Massachusetts, U.S.A. 02345.
Request for Information — Shorebird Color-marking
In 1979 the Canadian Wildlife Service will be
continuing a large-scale program of banding and
color-marking shorebirds in James Bay. Since 1974
over 38500 shorebirds have been captured, resulting
in more than 1700 ‘bird days’ of sightings of dyed birds
ranging from eastern Canada to South America.
Considerable valuable information on migration
routes and strategies is being obtained and observers
are again asked to watch for and report any color-
dyed or color-banded shorebirds. Reports should
include details of species (with age if possible), place,
date, color-marks and, if possible, notes on the
numbers of other shorebirds present. For color-dyed
birds, please record the color and area of the bird that
was dyed. For color bands and standard metal leg
bands, please record which leg the bands were on,
whether they were above or below the “knee,” the
colors involved (yellow or light blue), and the relative
position of the bands if more than one was on a leg
(e.g., right lower leg, blue over metal, etc.). All reports
will be acknowledged and should be sent to: Dr.
R. I. G. Morrison, Canadian Wildlife Service, 1725
Woodward Drive, Ottawa, Ontario, Canada
K1G 3Z7.
The Douglas Pimlott Scholarship in Environmental Studies
Innis College, University of Toronto, has estab-
lished a scholarship fund in memory of Professor
Douglas Pimlott, who for many years taught ecology,
resource management, and environmental studies at
the university. Among his many contributions, Dr.
Pimlott established an Environmental Studies pro-
gram, designed not for in-depth specialist training but
in the spirit of a liberal education concerned with the
environmental issues that face us all.
The scholarship will be awarded to an under-
graduate student who, like Doug Pimlott in his own
life, combines high academic achievement with dedi-
cated social involvement in environmental concerns.
It is hoped that sufficient funds can be raised to endow
the award.
Contributions should be made payable to the
Douglas Pimlott Fund, University of Toronto, and
sent to: The Principal’s Office, Innis College, 2 Sussex
Avenue, Toronto, Ontario MS5S 1J5.
i
Book Reviews
ZOOLOGY
Animals in Peril: a guide to the endangered species of Canada and the United States
By D.C. Grainger. 1978. MacMillan, Toronto. 192 pp.,
illus. $14.95.
This is another in a long line of books examining the
plight of animals in North America. The title is
misleading, however, because the book deals strictly
with mammals. Furthermore, I do not see this book as
“a guide to endangered animals of Canada and the
United States.” It is a slightly less than erudite review
of twenty mammals exploited by man at one time or
another since his arrival in North America from
Europe. In fact, some of the mammals considered are
no longer endangered, or have never appeared on an
endangered species list.
The mammals considered include the Black-tailed
Prairie Dog, Kaibab Squirrel, Spotted Bat, Fisher,
Pine Marten, Black-footed Ferret, Sea Otter, Wolver-
ine, Gray (timber) Wolf, Red Wolf, northern Kit Fox,
Mountain Lion (eastern cougar), Grizzly Bear, Wood
Bison, Muskox, Pronghorn Antelope, Florida Key
Deer, Big Horn Sheep, Walrus, and the northern
Manatee.
Grainger has geared his book for young readers or
novice biologists. His treatment of habits, habitat,
and life cycle biology is very rudimentary and often
suffers from too much brevity.
The saving grace of this book is the author’s
treatment of the cause and effect relationships
The Great Arc of the Wild Sheep
By James L. Clark. 1978. 4th Edition. University of Okla-
homa Press, Norman. 247 pp. Paper US $11.50.
“Many are inclined to think of the wild sheep as just
scattered here and there around the world with little
reason — but this is not so. Although they are rather
thinly dispersed, the sheep are confined to a relatively
narrow arc of mountain ranges which sweep across
three continents.” The Great Arc of the Wild Sheep
examines the wild sheep of the world.
“All wild sheep of the world may be divided into five
general groups: moufloniforms of the Mediterranean
Islands and Middle Eastern countries, Argalis or
Ammons of Central Asia, Asiatic Bighorns of Eastern
Siberia, Dall’s of Alaska and Northern British
Columbia, and North American Bighorns of western
North America.” Each species is treated separately
involved in the decline of some of these mammals. He
examines the role man has played in their decline and
documents efforts made to reverse the trend. For
example, by 1940 there were only thirty Florida Key
Deer left in existence, the result of overhunting.
Interest in this small deer on behalf of the U.S. Fish
and Wildlife Service, the Florida Game and Fresh
Water Fish Commission and many conservative
organizations, the National Audubon Society to
name one, was instrumental in preventing its extinc-
tion. Land was set aside as refuge and today there are
approximately 400 animals in existence. To this end,
Grainger leaves his readers with a glint of optimism.
Essentially, the book examines the history behind the
environmental manipulation of 20 mammals, and the
consequences of such action when poorly planned and
executed.
The text is accented with color portraits and black
line drawings of the mammals. A_ bibliography,
although broad in scope, is provided for those who
wish to continue on with the study of these mammals.
PAUL A. GRAY
Hough, Stansbury and Associates Ltd., Suite 409, 1265 East
Arthur Street, Thunder Bay, Ontario P7E 6E7
with each treatment generally containing information
on the species’ physical features, its habitat require-
ments, and its distribution.
Scattered throughout the text are stories of hunting
expeditions. Some of these hunting expeditions were
mounted in an effort to collect specimens for museum
collections; some merely hunting adventures.
The Great Arc of the Wild Sheep was first printed in
1964. This fouth edition does not include a great deal
of new information over the first edition.
To enjoy this book, one must be truly interested in
wild sheep.
PETER CROSKERY
Ontario Ministry of Natural Resources Ignace, Ontario POT
1TO
389
340 THE CANADIAN FIELD-NATURALIST
A Guide to the Birds of Venezuela
By Rodolphe Meyer de Schauensee and William H. Phelps,
Jr. 1978. Princeton University Press, Princeton, New
Jersey. xxii + 424 pp., illus. Cloth US $50.00; paper US
$19.95.
The publication of this guide is a noteworthy
addition to the ornithological literature of South
America in that it summarizes in one volume
descriptive, distributional, ecological, and taxonomic
information about the 1296 species of birds known to
have occurred in Venezuela.
The book comprises an introduction in which the
Venezuelan avifauna is summarized too briefly (114
page): the geographical and ecological features of the
country are dealt with briefly also (4 pages). Travel to
and in Venezuela is treated in a few words. A plan of
the book and short comments about the illustrations
conclude the section. A “Chart of a bird” and a “List
of families of Venezuelan birds” follow.
The scope of the book is limited to Venezuela but
nevertheless encompasses a very broad topic. The
book, to quote an excerpt from the introduction,
“contains a description of all the species of birds
known at present to inhabit Venezuela and its
adjacent islands ....” Most of those species, except for
some North American migrants, are depicted in the 53
plates assembled in the center of the book. A map of
Venezuela appears on both inside covers and cor-
responds to geographical entities mentioned in the
text.
Each family is introduced by a short paragraph in
which are given general characteristics of the family, a
short summary of the habitat preferences, other
ecological facts, and general information of interest.
The information dealing with each species is short
but factual. The authors have provided in addition to
the scientific name, Spanish and English common
names for all species. The latter are particularly well
chosen and descriptive. Descriptions are given in
telegraphic style and, to my knowledge, are accurate,
at least for the 25-30 species that I have checked
against specimens. Characteristic details peculiar to a
species are sometimes given in italics and are provided
for both sexes in species where sexual dimorphism is
present. Total length and occasional measurements of
certain body parts are provided (metric and imperial
systems). Descriptions appear to be adequate for a
large number of species but one wishes that more
distinguishing features had been given for similar
species. This would certainly be appreciated in the
field. Zoogeographical data are contained in a short
paragraph, while in addition to indicating the “life
zone” of the species, it summarizes the range in
Venezuela and in peripheral countries. The ecological
Vol. 93
and distributional data are necessarily brief but are
adequate in orienting the observer in the field. In
addition, this part often contains behavioral infor-
mation as well as descriptions of calls or call notes.
Line drawings of an exceptional quality by Michel
Kleinbaum supplement several descriptions. The
other illustrations are grouped in the center of the
book and “show virtually all of the resident species of
birds found in Venezuela.” Thirteen black-and-white
plates (i-xili) by Guy Tudor are of excellent quality
and are particularly efficient in depicting raptors in
flight.
The 40 color plates have been contributed by
Tudor, Wayne Trimm, John Gwynne, and Kathleen
Phelps. They are of very good quality in general but
Tudor’s are exceptional.
Field characters are given for each species opposite
each plate, or ina few cases, on following pages where
species are too numerous on a single plate. The layout
is attractive and species have been arranged in a
logical manner on the basis of similarities between
them; this is undoubtedly the most practical approach
to field identification. Of course, additional details on
distinguishing characters appear in the textual des-
criptions, which can be located rapidly, the page
number being indicated next to the species name. All
illustrations should make it easy for the field observer
to identify rapidly a large number of species, at least
the males. Flycatchers will no doubt continue to bea
nightmare!
A species list based on specimen records deals with
the birds of Isla de Aves on page 369. Subspecies
depicted in the illustrations are listed in an appendix,
followed by a short bibliography, which should have
more correctly been called “selected references.”
Indices to English and Spanish names, as well as to
genera and species are given in the last pages of the
book.
The cloth copy is not solidly bound and it appears
that it may not resist intensive use in the field.
Furthermore, the price of the cloth edition is very high
in comparison with the paper edition and I wonder
why hard covers alone cost $30.00 (USA)!
In spite of this shortcoming, this guide is the best
available to date for South America, and the authors
and artists are to be congratulated on their excellent
work. I highly recommend the book to anyone
interested in South American birds or planning to
watch birds in northern South America.
HENRI OQUELLET
National Museum of Natural Sciences, National Museums
of Canada, Ottawa, Ontario K1A 0M8
7)
1979
North American Moose
By Randolph L. Peterson. University of Toronto Press,
Toronto. 280 pp., illus. Paper $13.50.
This book, first published in 1955, has been re-
released in paperback. At the time the author did his
research it represented the only serious long-term
study on Moose in North America. It is still well worth
reading, both for the uninitiated and for those of us
actively working with this species. North American
Wild Geese
By M. A. Ogilvie. Buteo Books, Vermillion, South Dakota.
340 pp., illus. US $25.
Recently there have been several books written
dealing with waterfowl in a general manner (Johns-
gard 1975; Bellrose 1976; Palmer 1976). Unlike these
former works, “Wild Geese” deals only with geese,
and ona world scale as opposed to one continent.
The format is considerably different from the usual
and somewhat dreary collection of individual treat-
ments by species. Instead, subjects are examined
including classification, identification, ecology,
breeding, population dynamics, migration, distribu-
tion, and status. Within this framework, the various
species of geese are discussed. This tends to make the
text flow in a manner that is pleasant to read. Also
contributing to the ease with which this book may be
read is the author’s considerable talent in dealing with
often highly technical material in a fashion readily
understood by the amateur ornithologist. It is well-
illustrated, by Carol Ogilvie, with color plates; the
pen-and-ink sketches throughout are especially good.
The major criticism the reviewers had, of a general
nature, was the tendency towards too much detail on
certain subjects and limited coverage of others. For
example, ringing (banding) notably of Pinkfeet,
numbers in subpopulations, and molt migrations all
received extensive discussion. On the other hand,
behavior (other than during breeding), nutrition, and
physiology perhaps should have been covered in
greater detail. Bergmann’s rule is mentioned in several
locations. This rule is at best tenuous when applied to
geese, a fact noted by the author on p. 121. Ross’
Geese and Lesser Snow Geese do not illustrate the rule
as stated (p. 25). Over 65 percent of Lesser Snow
Geese breed south of most Ross’ Geese. This
discussion should have been limited or omitted
altogether.
The book contains many small errors. Map 25
should include heavily used areas at James Bay and
North Dakota. Snow Geese migrate directly to the
Gulf of Mexico from James Bay, not Hudson Bay.
BOOK REVIEWS 34]
Moose presents a foundation on which biologists have
built their knowledge of Moose. Asa follow-up to this
book and for a more recent compendium of the state-
of-the-art on Moose, I recommend Alces, Moose
Ecology, Les Presses de Université Laval, 1974.
IAN D. THOMPSON
P.O. Box 895, Cochrane, Ontario POL 1C0
Map 24 shows no migration out of areas 7 and 10
while some movement further south is known to
occur. Ross’ Geese, not using the west coast, winter
primarily in Texas as well as Louisiana (Palmer 1976).
Numbers of Canada Geese in the Mississippi Flyway
are over 700000 not 60000 (Hanson 1974). Cap
Tourmente has been used for closer to at least 500
than 400 years. North American waterfowl hunters do
not have to report their kills. The second reviewer was
surprised to see data quoted from his doctoral thesis,
yet not referenced (Prevett 1973). Minor errors such
as these can be expected in a book of this type,
particularly when a British author tackles the North
American waterfowl literature.
On the whole, this book is informative yet easy to
read. The author has done an excellent job of
compiling the extensive literature on geese into a
straightforward concise work. We recommend it
particularly for amateur ornithologists and biologists
not familiar with geese. Biologists will especially
appreciate the chapters on ecology and breeding.
References
Bellrose, F.C. 1976. Ducks, geese and swans of North ©
America. Harrisburg, Stackpole Books. 544 pp.
Hanson, R. 1974. Report on the December, 1973 inventory
of Canada Geese in the Mississippi Flyway States. U.S.
Fish and Wildlife Service Report. 13 pp.
Johnsgard, P. A. 1975. Waterfowl of North America.
Bloomington, Indiana University Press. 575 pp.
Palmer, R.S. (Editor). 1976. Handbook of North
American birds. Volume 2. New Haven, Yale University
Press. 521 pp.
Prevett, J. P. 1973. Family behaviour and age dependent
breeding biology of the blue goose, Anser caerulescens.
Ph.D. thesis, University of Western Ontario, London,
Ontario.
IAN D. THOMPSON! and J. P. PREVETT?
'P.O. Box 895, Cochrane, Ontario POL ICO
2Moosonee, Ontario POL 1Y0
342
Crows of the World
By Derek Goodwin. 1976. Cornell University
(Comstock), Ithaca, New York. 354 pp. US $28.50.
Press
It is fitting that Goodwin should contribute this
authoritative book on another of his favorite groups.
He began work on the jay about 30 years ago, and his
long tenure with the British Museum of Natural
History has allowed him further opportunities to
explore the fascinating crow family. His earlier work,
Pigeons and Doves of the World (1967), has recently
been reissued as a second edition (Cornell University
Press, 1977). The style, a welcomed combination of
scientific yet non-technical writing, is similar for the
two books. Because it summarizes much of what is
known and clearly shows what is unknown, it is an
excellent reference for ornithologists yet is very
readable for amateur naturalists.
The general format also is similar to his earlier
work. It begins with four general chapters on
nomenclature, adaptive radiation, plumage and
coloration, and behavior. The 41-page behavior
chapter discusses a myriad of behavioral characteris-
tics and is especially interesting.
Detailed species accounts are given in Chapter 5.
An introductory two-and-one-half pages of Chapter 5
provide an interesting account of man’s relationships
with corvids, touching upon their role in our folklore,
legend, and literature; our feelings toward their
predation on eggs and young of small birds; the crop
damages they cause; and their use as food. Where
species comprise a distinct group, an introductory
preface of taxonomic relationships and habits is
provided. Each of the 116 species is then covered in
detail — description, field identification, distribution
and habitat, feeding and general habits, nesting, voice,
display and social behavior, other names, and a list of
references. Length of the accounts varies from about
one-half page for the Banggai Crow, Flores Crow, and
Life of the Harp Seal
By Fred Bruemmer. 1977. Optimum (Prentice-Hall, Tor-
onto). 170 pp., illus. $25.
Fred Bruemmer has produced a very readable work
on the life of the Harp Seal. His style is pleasant and
flows easily as he provides accurate accounts of the
environment, culture, social structure, and history of
the Atlantic region where the sealing activity takes
place. His descriptions of the ice-environment are
especially good. His commentary on some aspects of
the biology of the Harp Sealare at times insightful and
competent. Sometimes, however, Bruemmer’s sci-
entific commentary is mixed with ethical considera-
THE CANADIAN FIELD-NATURALIST
Vol. 93
Sumatran Tree Crow to over eight pages for such well-
studied species as the jay and magpie. For several
species, there is no information provided other thana
description and range. The many “No information”
statements should certainly spur young ornithologists
to go “afield.” As the author states, one of the book’s
purposes is to indicate what is not known. About one-
fourth of the pages in the species accounts chapter
deal with the 16 species occurring in Canada and the
continental United States.
Robert Gillmore has done an excellent job
illustrating many of the species with line-drawings.
Three colored plates showing adults of 22 species and
juveniles of 4 species are nicely done and show the
variety of shapes, colors, and patterns within the
Corvidae. Range maps are provided for each species.
These are small, general, and do not give any
indication of seasonal occurrence. Migration or
movements, in general, is the only subject I feel is
slighted in the entire book. I would suspect that
because of their size, the maps may not be totally
accurate nor reflect up-to-date distributions. No
mention is made of the regular occurrence of the
Brown Jay insouth Texas. One other omission I noted
was the absence of a discussion of hybridization in
New World corvids, a rare but documented event.
Common and scientific names are indexed following
Chapter 5.
This fully referenced volume stands as an extremely
useful and important contribution. I believe that ali
readers will attest to the accuracy of the author’s
statement that the crow family is a numerous, diverse,
and successful group of passerine birds.
NOEL J. CUTRIGHT
Wisconsin Electric Power Company, 231 W. Michigan,
Milwaukee, Wisconsin 53201
tions which seem out of place. For example in a
discussion of the seal’s underwater hearing ability, he
points out that the seal is “as one with the sea,
... but when she must leave the sea... then she and her
kind are at the mercy of man.” The text is ripe with
anthropomorphic references to “sad and worried
mothers” and “jealous mates” which might be
forgiven as poetic licence. The cause of the seal is
taken up with such emotion, however, that the reader
is left to question whether Bruemmer is offering the
promised clear and unbiased account of the life of the
Harp Seal. Photographs are as important as the text
1979
of this coffee-table-size book. Ninety-eight color and
black-and-white photographs which range in size
from 2 X 3 inches to double-page spreads are con-
tained within the book’s 155 pages and many of them
are excellent, as is characteristic of the author’s other
books. But a seal pup, in various appealing poses,
appears in 61 of these pictures, as if one animal had
been photographed many times from different angles.
Most of the photographs are technically good, but
many are only fair. In the few underwater sequences
detail is difficult to resolve, and the large plate on page
101 is upside down.
Federal fisheries scientists will be disappointed with
Bruemmer’s half-page comment on Harp Seal
management. Most of it is cynical and disparaging,
and refers to the maximum sustainable yield as the
Holy Grail of management biologists. Bruemmer
spends much more time describing the “interfering”
activities of man (sealers, tourists, aircraft) in the
seal’s life.
Bruemmer’s description of sensory functioning in
Harp Seals includes an able discussion of much of the
data on the vision of seals and how it relates to their
behavior in air and under water. His descriptions of
audition and of vibrissae functioning, however, are
inaccurate and at times patently false. He states that
the seal’s audiogram in air is similar to, if not better
than, that of the human. In fact, though seals can hear
higher pitches than humans, they are at all fre-
quencies less sensitive to quiet sounds. His suggestion
that a seal could hear a man walking on the ice a mile
away is unlikely when one considers the relative
acoustic impedances of ice and water. As far as
whisker functioning is concerned, his statement that
seals use their vibrissae to detect changes in water
pressure, or the “tap of water against blocks of ice
pressed far beneath the floes,” and his implication that
the whiskers can perceive the turbulence of fish
movements over some distance are totally unfounded.
Current evidence indicates their vibrissae are not
sensitive enough for this.
His confident elaboration about the sense of smell
being the primary means by which mothers and pups
identify each other, and his contention that mothers
can recognize the call of their own pups is completely
unwarranted since such hypotheses have never been
tested. Similarly, to our knowledge, his statement that
pups are able to locate their mother’s teat by an
increase in skin temperature in the area of the nipple is
unsubstantiated. Also, he implies that the mothers
initiate weaning; however evidence from Harbor Seals
BOOK REVIEWS
343
shows that pups wean themselves.
Bruemmer describes in some detail the navigational
methods Harp Seals use to traverse the distance
between Greenland and their breeding grounds. In
fact, very little is understood about the way in which
any seal species plots its course over long distances. In
a similar vein he states without qualification that
Harp Seals can stay submerged for at least thirty
minutes and can dive to depths as great as 600 feet,
when in fact these numbers were obtained from the
Antarctic Weddell Seal.
In his discussion of the social behavior of Harp
Seals, Bruemmer asserts that these animals are
gregarious, and that during the breeding season the
female, when hauled out on the ice maintains a
“moveable territory” about herself that only her pup
can encroach upon. These are probably reasonable
hypotheses although they have not been directly
investigated. In further descriptions of reproductive
behavior, however, his portrayal of the events leading
to copulation are at best misleading and at worst
completely incorrect. He states that a single male
emerges the victor of an often bloody battle, to mate
on the ice with a willing female who has been wooed
by an hour-long display. In reality, nearly all
copulations occur in the water and therefore have
rarely been witnessed. The only published under-ice
observations of reproductive behavior in these ani-
mals suggest that the seals are probably promiscuous,
the females engaging in sexual displays with many
males. Furthermore, as is the case in most mammalian
species, true “bloody” battle is the exception rather
than the rule: agonistic behavior in this context has
become ritualized. Bruemmer’s description of an
hour-long display on the part of the “victorious” male
is the first such report.
For a decade Canadians have waited for an
independent and unbiased commentary on the Harp
Seal and its harvest. One might have expected:
Bruemmer to have provided it since he has produced a
number of good works. Readers hoping to find
enlightenment on this contentious issue will find this
book a disappointment.
TOM H. NORTHCOTT! and DEANE RENOUF?
‘Northland Associates Limited, P.O. Box 1734, St. John’s,
Newfoundland AIC 5P5
*Department of Psychology, Memorial University, St.
John’s, Newfoundland AIC 5S7
344 THE CANADIAN FIELD-NATURALIST
BOTANY
Edible Garden Weeds of Canada
By A. F.Szczawinski and N. J. Turner. 1978. National
Museums of Canada, Ottawa, Ontario. 184 pp. $8.95.
This book is the first of a four-book series on the
edible wild plants of Canada slated to be fully
published by 1980, under the auspices of the Canadian
National Museum of Natural Sciences. In this
context, Edible Garden Weeds gains a significance
that it would not otherwise have.
The volume is published in paper wrappers with a
wire binding that allows the book to lie flat open at
any page, a desirable feature in any cookbook. And in
fact, the great inspiration of this work is the
combination of the “standard” cookbook format with
the equally “standard” edible weeds or wild plants
format. In our experience it stands alone amongst
similar books on this count, other texts generally
immersing recipes in a chatty dialogue with no
comparable systematic layout of ingredients and
instructions.
Pages are odd sized (approximately 21 X 21 cm)
and, unfortunately, the use of space is wasteful; in fact
Edible Garden Weeds stands alone in this regard also.
The bulky size, and useless weight of many '4- to 7-
full pages certainly makes the book of less use to
“scouts, outdoorsmen, and others concerned with
survival.” We hope that this “spacious” format
approach is dropped in other books of this series by
the National Museum, for if it is carried through it will
be no less than a slap in the face for conservation.
The main body of discussion of species and recipes
is printed on white glossy paper with an attractive type
style and a rather nice, very dark green ink. All other
pages are of colored paper, brown in front and bright
yellow at the back. The many line drawings are clear
and crisp, and only rarely have the artists “fudged” the
smaller details of the plants. Several color photos add
a definite exciting touch to the book.
Actual contents of the text provide a good intro-
duction to the edible weeds dealt with; however, we feel
it is deficient and misleading in several specific and
general instances. The fact that many areas where
weeds grow prolifically have poisoned soil needs to be
emphasized and re-emphasized, especially to a domi-
nantly urban audience. This is a high paradox in this
book, for while people are warned away from a few
clearly edible species on rather slim grounds, no
mention is made of the dangers of lead toxicity
inherent in many an “unkempt boulevard.” The
plethora of dangerous industrial chemicals that today
contaminate many rail rights-of-way, roadsides, and
Vol. 93
vacant lots is likewise unmentioned. Asa general rule,
any weeds growing near urban or industrial develop-
ments should be avoided, but in fairness Edible
Garden Weeds is not alone in its lack of this warning.
Also, we wish that the nutritional virtues of weeds,
when growing in uncontaminated sites, had been far
more completely explored. Given the small number of
species and species groups treated (about 40),
wouldn’t it have been possible to produce a more
precise discussion of these virtues, and fill some of the
waste space in the book with, say, distribution/
abundance maps? Most of the text discussion is a
reasonably complete “standard” description of plants,
habitats, uses, etc., slightly modified better to fit the
Canadian context, and clearly organized under
headings.
The many recipes (about 130) without doubt
contain the most controversial aspects of this book.
They range from simple teas, salads, soups, and
spreads to lunches and even rather exotic meals. On
the one hand, the recipe layout and use of common
“supermarket” ingredients will be attractive to many
urban and suburban cooks. On the other hand,
natural food buffs can object to over forty ingredients
as chemical-ridden or simply “non-food” items. At
least fifteen of these could readily have more natural
or “real” foods substituted, and the complexity of
some recipes should be questioned. A certain balance
of nutrition and health seems to have been overlooked
in the attempt to make “weeds” credible to the general
cook.
This is not a book for the backwoods or farm, but
rather seems designed for suburban or estate resi-
dential environments and tastes. The attempt seems to
be to make “weeds” acceptable by associating them
with heavily flavored and often exotic ingredients
within a colorful, open and “artsy” publication style.
Much information is presented, but in no way can the
text be considered a critical analysis of the values and
uses of edible weeds and it stands on relatively equal
footing with several other easily available works. Any
one who wishes to understand properly the value of
edible weeds will still have to purchase a collection of
several books, and in this context the acquisition and
use of Edible Garden Weeds of Canada becomes a
matter of personal taste.
R.S.W. BOBBETTE! and LANDA L. KING?
131 Eccles St. N., Barrie, Ontario L4N 1Y1
28 Peel St., Apt. #3, Barrie, Ontario L4M 3K8
7s)
Ferns of the Ottawa District
By William J. Cody. 1978. Printing and Publishing, Supply
and Services Canada, Hull, Quebec KIA 0S9. 112 pp.
Soe:
The revised edition of this book is quite delightful.
There is a modern durable plastic binding with some
attractive silhouettes on it and the printer has done a
superb job with good modern clean type and the right
mix of various sizes of bold-face type, italicized type,
etc. By the clever use of dividing lines, the author has
solved the problem of associating the figures, the text,
and the distribution maps for each species. The
illustrations of selected herbarium specimens are by
and large the same as in the first edition of 1956, but
because they are on glossy paper, have slightly less
gray in the background and have a black line on three
sides of the plate, they seem infinitely clearer, sharper,
and more three dimensional. Another great im-
provement is the distribution maps, which are much
larger in size and so crystal clear that one feels one
could almost rush out to a given locality dot! The
keys are simple and workable and the addition of five
new species to the District (Pellaea atropurpurea, P.
glabella, Woodsia obtusa, W. oregana, and As-
plenium platyneuron) plus a plethora of new locality
dots, e.g., Ophioglossum from 5 to 21, brings the
revision right up to date. In these days of soaring book
costs, this book is a tremendous bargain and should be
purchased by anyone interested in ferns.
What are the shortcomings of this fine identifi-
cation manual? Most of my criticisms are a plea for
further information from authors. For example,
consider the distribution maps. Do the dots each
represent a herbarium specimen or do they represent
all records known to the author? Should a distribution
map represent the status of the species today or should
it include historical records (Dryopteris goldiana now
under a shopping mall)? What about sight records and
specimens in private herbaria?
Next the taxonomy, which might be dubbed
“authoritative conservative.” Admitting that the
names used are familiar ones and this manual is
probably not a good vehicle for radical taxonomy, it
still seems the author might include a statement of his
philosophy, or even a plaintive disclaimer that he is a
follower of, or influenced by, Gray’s Manual, Britton
and Brown, Wherry or whatever. It is most confusing
to the amateur to see Camptosorus in this book and
Asplenium in another; also Athyrium vs. Diplazium.
Similarly, the Long Beech Fern can be found in
Thelypteris or Phegopteris and as P. polypodiodes or
P. connectilis. Fortunately the author has supplied a
few synonyms for cross reference.
I do not like the treatment of Dryopteris, partly of
course because of my personal bias and knowledge of
BOOK REVIEWS
345
the genus. I feel that there should be six species treated
and not four and also wonder why D. X boottii is
singled out as the one hybrid recognized for the
district. The most common hybrid is alluded to on
p. 70 (D. X triploidea Wherry). The common name,
Florist’s Fern for var. spinulosa (not D. intermedia)
on p. 66 is most unfortunate. I cannot imagine a florist
using this fern and would deplore the attempt!
Another point concerns the designation of variety.
This seems to be a legacy of Gray’s Manual. Surely if
one 1s not going to recognize our Ostrich Fern as a
distinct species, it is at least a subspecies? Some
varieties seem a bit trivial or useless, e.g., Botrychium
matricariaefolium var. matricariaefolium, B. multi-
fidum var. intermedium, and Polystichum braunii
var. purshii, whereas others such as Dryopteris
cristata var. clintoniana are major biological species
as shown by cytogenetics. I think a few references for
the amateur who wishes to delve more deeply would
be extremely worthwhile.
Amateurs are also apt to ask for the rationale of the
sequence of genera as presented. Why is Dennstaedtia
between Polystichumand Athyrium? If no grouping is
intended, or no authority followed, then why not use
an alphabetical approach?
Ideally it would be nice if authors were a bit more
explicit about field characters or “tricks of the trade.”
How will you know a Pellaea when you see one,
instantaneously? For instance, emphasis on indusial
characters for Cystopteris and Woodsia when these
are obscure on mature fronds confuses amateurs. I
foresee difficulties for workers distinguishing between
Woodsia obtusa and Woodsia oregana with the key.
A minor annoyance in manuals is the rather
unspoken assumption that one can map a species for
all time if one just carefully “covers the ground.” This
is far too static a concept for plant populations.
Accordingly, when we are told under Dennstaedtia,
“Large ferns in colonies” and then later “perhaps it 1s
overlooked,” I would prefer the idea that it is
spreading and becoming more weedy in the district.
Similarly, with Asplenium platyneuron, | doubt it was
there until recently and will not be surprised if the
locality dots increase in numbers very shortly.
I feel the author has done a most commendable job.
What I am really saying is, give us more — more
references, more explanations, more taxonomic
philosophy, and we will gladly pay more!
DONALD M. BRITTON
Department of Botany and Genetics, University of Guelph,
Guelph, Ontario NIG 2WI
346
THE CANADIAN FIELD-NATURALIST
Vol. 93
Dyes from Lichens and Plants: a Canadian dyer’s guide
By Judith Waldner McGrath. 1977. Van Nostrand Reinhold,
Toronto. 144 pp., illus. $14.95. Translated into Inutituut
and syllabics.
How this book came to be written is as interesting as
the information on dyes from native plants. Mrs.
McGrath is a dyer, a weaver, a sculptor, and a teacher.
In recent years she has lived in several northern
settlements following her husband in his work with
the Economic Development Council.
Spence’s Bay, where the book was written, is a
community of 400 people on the Boothia Peninsula, a
tiny speck on the bleak arctic landscape when seen
from the air. On arrival, it was a shock to find native
people in modern clothes and living in ordinary
homes, but as Mrs. McGrath made friends with the
Inuit she recognized that cultural differences do exist.
In the brief arctic summer flowers bloom every-
where. Continual daylight encourages growth but
high winds, lack of moisture, and occasional snow
cause severe stress to the environment. Yet close to the
ground, where the air is warmer, grow some 900
species of flowering plants and perhaps 1000 lichens.
It was here that Mrs. McGrath began her search for
dye stuffs. Each find was shared with her new friends.
Soon an experimental workshop was set up, opened
throughout the long winter night so that women could
work when it was convenient. The following summer a
grant made it possible to gather plants farther afield
and to experiment with colors that could be obtained
from them. To the Inuit women, used to bright
commercial dyes, the new muted shades were exciting.
Designs made from them won awards, and official
parkas for the 1974 winter games were designed and
made here. By 1977, when the McGraths left, the craft
shop was becoming a co-operative — 32 people were
involved in the new cottage industry.
Much of the book is devoted to the plants them-
selves, to the parts used and the amount of dye
material needed for best results. It is impressive that
from the beginning there was concern for the ease with
which the fragile ecological balance could be destroy-
ed. Spot collecting was done over a wide area. Only a
part of each plant was taken, leaving some roots for
propagation and some flowers to set seeds. Material
was preserved by drying, by fast freezing of plants, or
by preparing and freezing the dye bath. Inall, 17 basic
dye methods or recipes were evolved. Of these nine are
for lichens, the rest for flowering plants, seaweeds,
and mosses.
Few colors remain fast on fiber without the use ofa
mordant such as alum, iron nails, or copper.
Mordants combine chemically with the dye and the
fabric to fix the color. Variation in shade can be
obtained by changing the mordant used or the time at
which it is added. For the beginner the easiest plants
are those with their own mordant such as sumac,
which contains tannin, and the lichens which have
their own acids.
There are chapters on the people and their crafts
and on the land itself, brought to life by black-and-
white photographs. Beautiful color plates illustrate
groups of plants and display small tie samples of wool
colors obtained from them. There are lists of useful
equipment, of stores that carry supplies for dyers, an
index of plants by colors they can produce, another of
plants in more temperate parts of Canada that will
produce comparable colors, and one giving botanical
as well as common names. In preparing the book,
government departments in both Ottawa and the
Northwest Territories were consulted. Identification
of all plants and lichens was confirmed at the National
Herbarium.
My only complaint concerns the color plates. They
are very attractively arranged and colors are good, but
there is no explanation of the photographs. In this
section, one finds only outlines of illustrations with
difficult-to-decipher numbers and letters. These illus-
trations would be so much more useful in the part of
the book where plants and dye colors are described.
Except for the difficulty with color plates the
material is well and interestingly prepared. The author
knows her subject and has been generous with that
knowledge. Mrs. McGrath says that care in each step
from washing of the fiber to mordanting and dyeing
affects how well and evenly colors take. Don’t worry
about the experts: read them and listen to them, but
the only sensible course is to experiment yourself.
Although the book is presumably intended for
weavers and dyers, many people unfamiliar with the
craft will find pleasure in it. The book is sturdily
bound with an eye-catching jacket. It was written and
published in Canada. Highly recommended.
MARY I. MOORE
Petawawa Forest Experiment Station, Canadian Forestry
Service, Chalk River, Ontario KOJ 1J0
1979
ENVIRONMENT
Human Activity and the Environment
By Statistics Canada. 1978. Statistics Canada, Ottawa.
190 pp., illus. Paper $2.80 in Canada; $3.40 elsewhere.
Statistics Canada has attempted to put together a
“statistical series detailing human activities which
have a potential for imposing stress on the natural
environment.” The resultant book has seven chapters
which are entitled watersheds; agriculture; forests;
fisheries; transportation; manufacturing; energy. It is
an interesting and welcome initiative by Statistics
Canada.
A novel section contains the presentation of certain
social economic data by the natural physiographic
unit of the watershed. All of Canada is delineated
along the boundaries of the major, and some minor,
watersheds. Some of the data given by these units
include human population; watershed area; human
population density; farms and farmlands fertilized,
sprayed and irrigated; livestock numbers; and
industrial activity.
Some interesting examples of data from the text are
given below. The large increase in the sales of pest
control products, which are also known to some of us
as biocides, over the last 30 years is somewhat
disturbing. Since 1971 the increase appears to take the
shape of an exponential curve! Statistics Canada
outlines the striking decline in the weight of fish taken
by Canadians off Canada’s shores from 1968 onward.
The corresponding increase in landings by other
nations, most notably the Soviet Union, provokes
thought.
An attempt is made to make the text visually
interesting. This attempt works reasonably well. One
illustration which stands out is the complex map of
noise contours around Toronto International Air-
port. The noise magnitudes are presented by various
shades of blue. The many people that live in the area
BOOK REVIEWS
347
and are subject to the noise disturbances often see red
on this issue.
The data presented is scattered in content and given
uneven treatment. Nevertheless one must sympathize
with the authors’ considerable task of choosing
subjects in such a broad field.
There are some obvious biases ina document of this
kind. First, the environmental phenomena must be
easily measurable and quantifiable. Second, there
must be an existing measurement program underway.
Most readers of this journal could easily think of
many critical environmental issues that badly need
methodological measurement. For example, human
activities often put considerable stress on wildlife
populations but usually we have only partial data on
what population changes do occur.
Some classes of data that might be given
consideration for future editions include the follow-
ing:
1) The attitudes of Canadians towards the environ-
ment. Who cares? How much? About what?
2) The recreational pursuits of Canadians in terms of
dollars and time spent.
3) Canadian land-use changes over time. The existing
section in the book should be expanded.
4) Air quality data.
5) Water quality data.
6) Various wildlife populations data.
It must be kept in mind that this first attempt has
broken considerable new ground. Hopefully, refine-
ment and enlargement will take place with future
editions.
PAUL F. J. EAGLES
School of Urban and Regional Planning, University of .
Waterloo, Waterloo, Ontario N2L 3Cl
Vermilion Lakes Banff National Park: an introductory study
By the Bow Valley Naturalists. 1978. Bow Valley Natural-
ists, Banff, Alberta. 68 pp., illus. Paper $3.00.
The Vermilion Lakes area, immediately west of the
town of Banff, in Banff National Park, has long beena
focus of interest for visiting and resident naturalists.
The lakes themselves are large, shallow ponds which
(for the mountains) are highly productive aquatic
systems. The view of the north end of Mount Rundle
across the lakes is a famous and frequently photo-
graphed sight and has become almost synonymous
with Banff.
The richness of the area, coupled with increased use
by visitors and the threat of further impact by highway
construction along the northern edge of the lakes has
heightened concern for its future. This publication is,
in part, an attempt to increase public awareness of the
values of the Vermilion Lakes in the face of new
pressures on them.
The study covers a range of cultural and natural
history fields, each treated in a separate section and
based on the field studies and research carried out by
various members of the Bow Valley Naturalist Club in
348
the mid-1970s. A set of recommendations regarding
the future use and development constraints on the
lakes are made which result from an assessment of the
data assembled during the study.
All of this is pulled together in a tight, attractive,
paper-covered booklet which ts effectively illustrated
by line drawings, photographs, and maps. A some-
what cumbersome poem about the lakes starts it all
off, and the booklet ends with a fairly extensive list of
references. It is well put together, with clear easily
readable type, crisp appropriate photographs and
some excellent pen-and-ink drawings. In format and
production it is a pleasing and effective document to
deal with.
The study is intended to introduce the reader to the
Vermilion Lakes. It does that quite admirably. |
suppose my only major criticism is that the authors
held themselves to this. Detailed flora and fauna data
are not provided despite quite extensive studies in
some of these fields. Appendices are maintained by
the Bow Valley Naturalist which contain raw lists of
the observations reported during the study period, but
these are not easily accessible. In the booklet proper,
only very general statements of status are included
(e.g., “occasional in wet spruce forest” or “fairly
common. Nests”). This does introduce the reader to
the subject; it would have given him so much more if
that extra effort had been made to pull these obser-
vational data together into precise, clearly docu-
mented discussions. It seems that making such
statements as ... “The Cougar appears to be on the
verge of extinction in Banff National Park,” and the
American Bittern ... “Must nest but no records,” are
quite inadequate without more supportive documen-
tation and discussion.
MISCELLANEOUS
Weather Almanac
Edited by J. A. Ruffner and F. E. Bair. 1977. 2nd edition.
Gales, Detroit. 728 pp. US $25.
The Weather Almanac is useful to both the layman
who is interested in weather phenomena and the
technical people who use weather data in their jobs.
The section on “Weather Fundamentals” would be
a better section to put first than the “Atlas Format
Weather.” I recommend that this section be read first
to both interest and inform the reader.
I am slightly disappointed that the 1941-1970
tridecade was used for most tables when an upgrade
using data from 1941-1976 is easily available from the
THE CANADIAN FIELD-NATURALIST
Vol. 93
A volunteer effort of this nature is very dependent
on the quality and quantity of the individual effort
contributed by concerned naturalists. Inevitably, the
quality of content varies throughout in such an effort.
The herpetological section, for example, is very
strong, whereas the botanical material is not so. I
understand that this latter area suffered considerably
because of Parks Canada’s unwillingness to provide a
collecting permit for the study!
Although quite free of typographical errors, a
number of spelling errors and omissions do occur
(e.g., Gallium for Galium, incomplete scientific name
for Alder Flycatcher, Spirea for Spiraea). These are
infrequent enough, however, neither to affect seri-
ously the credibility of the document nor greatly to
impair its readability.
The Vermilion Lakes study is a fine effort by a
number of concerned and dedicated naturalists who
have obviously gone to considerable personal expense
and effort to increase our awareness and appreciation
of an important natural area in Banff National Park. I
hope it inspires other groups to embark on similar
ventures and I would certainly recommend it to those
readers who would like to know more about Banff. I
recommend it even more strongly to those of us who
often feel that individuals and small groups of
naturalists can’t do much to document and publicize
the values of an area which we cherish. The Bow
Valley Naturalists have shown that it can be done and
they are to be congratulated for their fine efforts.
D. F. BRUNTON
Kananaskis Provincial Park, Alberta Department of
Recreation and Parks, Canmore, Alberta TOL 0MO
weather service. This is especially important since new
extremes were set in 1976 and 1977 and the almanac
does not, therefore, explain the harsh winters for
example.
The “Glossary of Weather Terms” is certainly
useful as it explains in general terms most of the
meteorologist’s jargon.
The section on “Climates of the World” is very
useful for travellers but lacks an overview of
“Climate” in general. It would be better to put our
present world climate into an historical perspective. We
are presently in an abnormal period of climatic
1979
history. We are coming to the end of an interglacial
warm period and the tridecade data reflects only this
period. Throughout most of the world’s existence, the
climate has been “glacial” and the fact that it is not
now is both interesting and gives man a sense of the
enormous impact of climate. This leads naturally into
the “Health and Weather” section and the awareness
that there are scientific relationships between health
and the weather.
The descriptions and safety rules for “storms and
severe weather” are clear and especially useful for self-
protection in the winter. Folklore (such as p. 119) and
rules of thumb (distance to storms, be your own
forecaster) add immeasurably to the dry textual
information. Earthquakes are not strictly speaking a
weather phenomenon, but the tables do allow the
reader in a specific area to review this phenomenon
both historically and currently.
The section on “Air Pollution” and its effects (Table
5.2) is slightly misleading in that there is conflicting
evidence on effects and levels of pollution at which
they occur. Similarly Figure 5.14 does not cover the
case of a plume that is hot enough to break through a
weak inversion, thus trapping pollutants above it. In
this case almost none of the pollutant comes down to
the ground to affect receptors.
Introduction to Mathematics for Life Scientists
By E. Batschelet. 1976. 2nd edition. Springer-Verlag,
New York. 643 pp. Cloth US $27.60; paper US $11.95.
Most frequently, book reviewers possess some ex-
pertise in the subject area of the book they are
reviewing. This review departs from that tradition.
My mathematics is best described as shakey. For
precisely this reason I feel qualified to recommend
Introduction to Mathematics for Life Scientists as a
study guide and reference book for those who feel
uncomfortable with the steadily increasing number of
non-linear functions, differential equations, and ma-
trix operations to be encountered in the current
literature.
To facilitate study the chapters are comprised of
bite-sized sections with alternating theory and ex-
amples or applications. Most examples are from the
recent literature and referencing i$ thorough through-
out. Each chapter is concluded by a section of
problems, graded in difficulty, with answers for odd-
numbered ones.
Most commendably the author starts at the
beginning with a review of basic algebra, a knowledge
of which is all too often erroneously assumed by other
BOOK REVIEWS
349
The section on “Energy and Weather” is an
excellent introduction to the fundamentals of energy
conservation for the days of restraint ahead.
The “Weather of Selected U.S. Cities” is a good
general summary of climate information and is well
documented and presented. The “Atlas Format
Temperatures,” on the other hand, is confusing and,
because of the type size, almost unreadable. It would
be more useful to have one map showing the climate
station locations and tables of highs, lows, and
averages. The “normal monthly precipitation and
snowfall” maps give easy reference to the amount of
rain and snow visually but the type is too small to read
for quantitative information.
Although there are a few typographical errors in the
text, on the whole the almanac would be a very useful
addition to either a layman or weather scientist’s
library.
JAMES W.S. YOUNG
Atmospheric Environment Service, 4905 Dufferin Street,
Downsview, Ontario M3H 514
texts of similar intent. Subsequently, linear, power,
periodic, and logarithmic and exponential functions
are dealt with. Chapters on ‘limits’ and ‘differential
and integral calculus’ build up to a treatment of
differential equations. Chapters on graphical methods
and probability are essentially self-contained. On the
other hand the chapter ‘matrices and vectors’ requires
considerable familiarity with earlier sections of the
book.
A detailed list of contents and a thorough index
make this a book to which one can refer easily. The
clear concise text constantly refers back to the precise
place where a difficult concept or step, likely to be
troublesome, was treated.
My own fruitful experiences with this book have led
me to the opinion that it provides an excellent starting
point for any biologist who feels restricted by lack of
mathematical expertise.
ROBIN MAHON
Department of Zoology, University of Guelph, Guelph,
Ontario NIG 2W1
350
NEW TITLES
Zoology
Adventure of the stone man. 1978. By F. Hamerstrom.
Illustrated by W. Kimber. F. Hamerstrom, Plainfield,
Wisconsin. US $6.95 plus 50¢ postage.
Alaska whales and whaling. 1978. Alaska Geographic
Volume 5, Number 4. Alaska Northwest, Anchorage.
144 pp., illus. US $11.95.
Animals in danger. 1979. By Joseph Lucas and Susan
Hayes. Clarke Irwin, Toronto. Paper $5.95.
*Biology of fishes. 1979. By Carl E. Bond. Saunders,
Toronto. 514 pp., illus. $22.75.
Biology of insect eggs: Volume I, general biology; Volume II,
the eggs of insect families; Volume III, indexes and biblio-
graphy. 1979. Pergamon, Elmsford, New York. cl500 pp.,
US $330.
Biology of Reptilia: Volume 7, ecology and behavior. 1977.
Edited by A. C. Gans and D. W. Tinkle. Academic Press,
New York. xvi + 720 pp., illus. US $60.60.
{Birds of the Gainsborough-Lyleton Region (Saskatchewan
and Manitoba). 1979. By Richard W. Knapton. Special
Publication Number 10. Saskatchewan Natural History
Society, Regina. 72 pp. Paper $3.
Bird sounds and their meaning. 1977. By R. Jellis. British
Broadcasting Corporation. 256 pp., illus. US $11.
The Bobcat of North America. 1978. By S. P. Young.
University of Nebraska Press, Lincoln. xv + 193 pp. Cloth
US $11.50; paper US $3.50.
tCamel quest. 1978. By Anne Innis Dagg. York Press,
Toronto. 150 pp., illus.
Contrasts in behavior: adaptations in aquatic and terrestrial
environments. 1979. Edited by E.S. Reese and F. J.
Lighter. Wiley-Interscience, New York. 406 pp. US $27.50.
Field guide to the seashells of the world. 1978. By G.
Linder. Translated from 1975 German edition by G. Vevers.
Van Nostrand Reinhold, New York. 272 pp., illus. Cloth
US $12.95; paper US $8.95.
tFossils of Ontario: Part 1, the trilobites. 1979. By Rolf
Ludvigsen. Royal Ontario Museum, Toronto. 96 pp., illus.
$7.50.
+General patterns of invertebrate development. 1975. By
Gary J. Brusca. Mad River Press, Eureka, California.
134 pp., illus.
TA guide to the jellyfish of Canadian Atlantic waters/ Guide
des méduses des eaux canadiennes. 1977. By C. T. Shih.
Natural History Series de l’Atlantique. Number 5. National
THE CANADIAN FIELD-NATURALIST
Vol. 93
Museums of Canada, Ottawa. 90 pp. English + 90 pp.
French, illus. Free.
Handbook of common New Guinea beetles. 1977. By J. L.
Gressitt and R. W. Hornabrook. Wau Ecology Institute
Handbook No. 2. Bishop Museum Press, Honolulu. vin +
88 pp., illus. Paper US $4.50.
The history and breeding biology of Canada Geese of
Marshy Point, Manitoba. 1978. By James A. Cooper.
Wildlife Monographs No. 61. Wildlife Society, Washington.
87 pp. US $3.15.
Insect clocks. 1979. Edited by D. S. Saunders. Pergamon,
New York. 288 pp., illus. US $11.
{The insects and arachnids of Canada: Part 5, the crab spiders
of Canada and Alaska (Araneae: Philodromidae and
Thomisidae). 1978. By Charles D. Dondale and James H.
Redner. Canada Department of Agriculture Publication
1663. Supply and Services Canada, Hull. 255 pp., illus. $7.50
in Canada; $9 elsewhere.
} Love affair with a Cougar. 1978. By Lyn Hancock. Double-
day, Toronto. 252 pp., illus. $9.95.
Mayflies of Michigan trout streams. No date. By J. W. and
F. A. Leonard. Bulletin 43. Reprinted. Cranbrook Institute
of Science, Bloomington Hills, Michigan. illus. $6.95.
Mexican wilderness and wildlife. 1978. By B. Tinker.
University of Texas Press, Austin. x11 + 131 pp. US $9.95.
Muskrats and marsh management. 1978. By P. L. Erring-
ton. University of Nebraska Press, Lincoln. 183 pp. Cloth
US $10.95; paper US $3.25.
Origins: what new discoveries reveal about the emergence of
our species. 1979. By Richard Leakey and Roger Lewin.
Dutton (Canadian distributor Clarke Irwin, Toronto). Cloth
$29.50; paper $11.95.
Patterns of primary production in the biosphere. 1978.
Edited by Helmut F. H. Lieth. Academic Press, New York.
368 pp. US $27.50.
Planning for wildlife in cities and suburbs. 1978. By D. L.
Leedy, R. M. Maestro, and T. M. Franklin. U.S. Fish and
Wildlife Service, Washington. 64 pp. Paper US $2.50.
Populations of small mammals under natural conditions.
1978. Edited by D. P. Snyder. Papers from a symposium,
Linesville, Pennsylvania, May 1976. University of Pitts-
burgh Pymatuning Laboratory of Ecology, Linesville,
Pennsylvania. xiv + 238 pp., illus. US $8.50.
Social insects Volume 1. 1978. Edited by Henry R. Her-
mann. Academic Press, New York. 456 pp.
1979
{Standard common and current scientific names for North
American amphibians and reptiles. 1978. By J. T. Collins,
J. E. Huheey, J. L. Knight, and H. M. Smith. Miscellaneous
Publications, Herpetological Circular No. 7. Society for the
Study of Amphibians and Reptiles, Lawrence, Kansas.
36 pp. Paper US $2.50. Order from Douglas H. Taylor,
Department of Zoology, Miami University, Oxford, Ohio
45056.
Statistical inference from capture data on closed animal
populations. 1978. By D. L. Otis, K. P. Burnham, G. C.
White, and D. R. Anderson. Wildlife Monographs No. 62.
Wildlife Society, Washington. 135 pp. US $3.50.
Strategies in cold. Natural torpidity and thermogenesis.
1978. Edited by Lawrence C. H. Wangand Jack W. Hudson.
Academic Press, New York. 734 pp. US $33.
Texas wildlife. 1978. By D. Baxter, T. L. Clark, and J.
Jefferson. Texas A & M University Press, College Station,
Texas. 196 pp. US $22.50.
Wildlife management in wildernesses. 1978. By C. A.
Schoenfeld and‘ J.C. Hendee. Boxwood, Pacific Grove,
California. 172 pp. Paper US $3.95.
Botany
The biology and chemistry of the Compositae. 1977.
Edited by V.H. Heywood, J.B. Harborne, and B. L.
Turner. Papers from a symposium, Reading, England, July
- 1975. Academic Press, New York. 1190 pp., illus (2 volumes).
US $107.50.
A checklist of names for 3000 vascular plants of economic
importance. 1977. By E.E. Terrele. Superintendent of
Documents, Washington. 11 + 202 pp. Paper US $3.50 + 25%
foreign handling.
Checklist of vascular plants of the Ottawa-Hull Region,
Canada/ Liste des plantes vascularies de la région d’Ottawa-
Hull, Canada. 1978. By John M. Gillett and David J.
White. National Museums of Canada, Ottawa. 155 pp.
Color in plants and flowers. 1978. By J. and S. Proctor.
Everest, New York. 116 pp., illus. US $9.95.
Embryology of gymnosperms. 1978. By Hardev Singh.
Borntraeger, Berlin. 302 pp., illus. DM 138.
Essays in plant taxonomy. 1978. Edited by H. E. Street.
Academic Press, New York. xxiv + 304 pp., illus. US $24.50.
Ferns of north-western Himalayas. 1979. By K. K. Dhir.
Cramer, Lohmar, Germany. cl60 pp., illus. cDM 40.
Fruits of angiosperms. 1977. By Ingrid Roth. Borntraeger,
Berlin. 675 pp., illus. DM 288.
+How to identify mushrooms to genus. I: macroscopic
features. 1973. By David L. Largent. Mad River Press,
Eureka, California. 86 pp., illus.
BOOK REVIEWS
3)3)II
+ How to identify mushrooms to genus. II: field identification.
1977. By David L. Largent and Harry D. Thiers. Mad River
Press, Eureka, California. 32 pp.
+How to identify mushrooms to genus. III: microscopic
features. 1977. By David L. Largent, David Johnson and
Roy Watling. Mad River Press, Eureka, California. 148 pp.,
illus.
+How to identify mushrooms to genus. IV: keys to families
and genera. 1977. By Daniel E. Stuntz. Mad River Press,
Eureka, California. 94 pp.
*Identifying grasses: data methods and illustrations. 1977.
By H. T. Clifford and L. Watson. University of Queensland
Press (Distributed by Prentice-Hall, Herts, England).
146 pp. £ 11.20.
Introduction to world vegetation. 1978. By A. S. Collin-
son. Allen and Unwin, Boston. 202 pp., illus. Cloth
US $16.25; paper US $8.75.
+Lehrbuch der Pflanzenphysiologie: 3, vollig Neubearbeitete
und erweiterte Auflage. 1978. By H. Mohrand P. Schopfer.
Springer-Verlag, Berlin. 639 pp., illus. DM 78.
A monograph of the genus Jubula Dumortier. 1978. By
W.R. Guerke. Cramer, Lohmar, Germany. 144 pp., illus.
DM 40.
The mosses of arctic Alaska. 1978. By W.C. Steere.
Cramer, Lohmar, Germany. 508 pp., illus. DM 150.
Nuclear division in the fungi. 1978. Edited by I. Brent
Health. Academic Press, New York. 256 pp. US $16.
The pine tree book. 1979. By Russel Peterson. Illustrated
by Patricia Wynne. Based on the Arthur Ross Pinetum, New
York. Brandywine (Canadian distributor Clarke Irwin,
Toronto). Cloth $19.50; paper $10.50.
Plant cold hardiness and freezing stress. 1978. Edited by
P. H. Liand A. Sakai. Academic Press, New York. 428 pp. -
US $21.50.
Plant life in anaerobic environments. 1978. Edited by D. D.
Hook and R. M. M. Crawford. Ann Arbor Science, Ann
Arbor, Michigan. x + 564 pp., illus. US $28.
The pteridophytes of Kansas, Nebraska, South Dakota and
North Dakota, U.S.A. 1979. By A. J. Ott-Petrik. Cramer,
Lehr, Germany. 350 pp., illus. cDM 80.
Recognition of tree species on aerial photographs. 1978. By
L. Sayn-Wittgenstein. Forest Management Institute In-
formation Report FMR-X-118. Environment Canada,
Ottawa. 97 pp., illus. Free.
The species problem in Cannabis: science and semantics.
1979. Ernest Small. 2 Volumes. Corpus, Toronto. 224 and
160 pp. Cloth $16.95 and $14.95 or $28 set; paper $10.95 and
$9.95 or $18 set.
352
A taxonomic monograph of Equisetum subgenus Equise-
tum. 1978. By R. L. Hauke. Cramer, Lohmar, Germany.
72 pp., illus. DM 40.
*Vascular plant families. 1977. By James Payne Smith, Jr.
Illustrations by Katheryn E. Simpson. Mad River Press,
Eureka, California. 320 pp., illus. US $7.85.
Water deficits and plant growth. 1978. Edited by T.T.
Kozlowski. Academic Press, New York. 320 pp. US $31.50.
Environment
tAnalysis of ecological systems. 1979. Edited by David J.
Horn, Rodger D. Mitchell, and Gordon R. Stairs. Ohio
State University Press, Columbus. 1x +312 pp., illus.
US $27.50.
Applications of ecological (biophysical) land classification in
Canada. 1979. Edited by C. D. A. Rubec. Proceedings of
the Second Meeting Canada Committee on Ecological
(Biophysical) Land Classification, 4-7 April 1978, Victoria.
Ecological Land Classification Series Number 7. Lands
Directorate, Environment Canada. Supply and Services
Canada, Hull.. xxix + 396 pp., illus. $7. in Canada; $8.40
elsewhere.
Biogeography and adaptation: patterns of marine life. 1978.
By G. J. Vermeij. Harvard Press, Cambridge, Massachu-
setts. xvi + 332 pp., illus. US $25.
+ Catalogue of type invertebrate, plant, and trace fossils in the
Royal Ontario Museum. 1978. By J. Waddington, P. H.
von Bitter, and D. Collins. Life Sciences Miscellaneous
Publications. Royal Ontario Museum, Toronto. 151 pp.
Paper $7.50.
+Contrasts in behavior. 1979. Edited by Ernst S. Reese and
Frederick J. Lighter. Wiley, Somerset, New Jersey. x1 +
406 pp., illus. US $27.50.
The dark range: a naturalist’s night notebook. 1978. By
David Rains Wallace. Sierra Club, San Francisco. xii +
132 pp., illus. Cloth US $15; paper US $8.95.
Discovery processes in modern biology. 1979. By W.R.
Klemm. Krieger, New York. US $14.95.
The economics of environmental quality. 1978. By E. S.
Mills. Norton, New York. 304 pp., illus. US $11.95.
}Energy, economic and ecological relationships for Gotland,
Sweden: a regional systems study. 1978. By A. M. Jansson
and J. Zucchetto. Ecological Bulletins 28. NFR, Stockholm.
152 pp., illus. Paper SwCr 70 (c$16).
Fungi, man and his environment. 1978. By H. J. Brodie.
University of Toronto Press, Toronto. xii + 132 pp., illus.
$10.
Georgian Bay Islands National Park integrated resource
survey. 1978. By R. Hirvonen and R. A. Woods. Forest
THE CANADIAN FIELD-NATURALIST
Vol. 93
Management Institute Information Report FMR-X-117.
Environment.Canada, Ottawa. vill + 82 pp. + appendices,
illus. Free.
Joy of nature: how to observe and appreciate the great
outdoors. 1979. By Reader’s Digest, Montreal. 352 pp.,
illus. $16.96.
{Muskeg and the northern environment in Canada. 1977.
Edited by N. W. Radforth and C. O. Brawner. By the
Muskeg Committee, NRC Associate Committee on Geo-
technical Research. University of Toronto Press, Toronto.
399 pp., illus. $35.
The natural history of the land of the bible. 1978. By A.
Alon. Doubleday, New York. 276 pp. US $12.95.
{Naturalist’s seashore guide: common marine life of the
northern California coast and adjacent shores. 1978. By
G. J. Brusca and R. C. Brusca. Mad River Press, Eureka,
California. 215 pp., illus. US $8.95.
*Nature detective. 1978. By Hugh Falkus. Gollancz (Cana-
dian distributor Clarke Irwin, Toronto). 256 pp., illus.
$22.50.
Nature’s oddballs. 1978. By L. Zappler, Doubleday, New
York. vill + 148 pp. US $5.95.
Oil, ice and climate change: the Beaufort Sea and the search
for oil. 1978. By Allen Milne. Edited by R. J. Childerhose.
Institute of Ocean Sciences, Sidney, British Columbia.
103 pp., illus. Paper $2.50.
Operating manual for spaceship earth. 1978. By R. B.
Fuller. Dutton, New York. 143 pp. US $2.95.
Pond littoral ecosystems. 1978. Edited by A. Dykyjova and
J. Dvet. Springer-Verlag, New York. 480 pp. US $49.
Population regulation. 1978. Edited by R.H. Tamarin.
Benchmark Papers in Ecology, Volume 7. Dowden, Hutchin-
son and Ross, Stroudsburg, Pennsylvania. xvi + 390 pp.,
illus. US $27.50.
St. Lawrence Islands National Park and surrounding areas
integrated resource survey. 1978. By R. Hirvonen and R.
Woods. Forest Management Institute Information Report
FMR-X-114. Environment Canada, Ottawa. viii + 57 pp. +
appendices, illus. Free.
Miscellaneous
The aurora borealis. 1979. By S. I. Akasofu. Alaska Geo-
graphic Volume 6, Number 2. Alaska Northwest, Anchor-
age. 96 pp., illus. US $9.50.
Beyond the green revolution: the ecology and politics of
global agricultural development. 1979. By Kenneth A.
Dahlberg. Plenum, New York. c215 pp. US $17.95 + 20% in
Canada.
1979
Camping and back packing: a guide to information sources.
1979. Edited by Cecil F. Clotfelter and Mary Clotfelter.
Gale, Detroit. c350 pp. US $22.
Diary of John Clifford Higgins, 1906-1914: a diary of
observations on birds. 1978. Edited by William W. Judd.
Phelps, London, Ontario. 68 pp. $5.
Early naturalists and natural history societies of London,
Ontario. 1979. By Wiliam W. Judd. Phelps, London.
216 pp., illus. $7.50.
Energy from the waves. 1979. By D. Ross. Pergamon,
Elmsford, New York. 80 pp., illus. Cloth US $15; paper US
$7.50.
Hierarchical analysis of water resources systems: modeling
and optimization of large scale systems. 1977. By Y. Y.
Haimes. McGraw-Hill, Hightstown, New Jersey. 478 pp.
US $39.50.
Introduction to marine pollution control. 1979. By Jerome
Williams. Wiley-Interscience, Somerset, New Jersey.
c256 pp. c US $30.
*Introduction to mathematics for life scientists. 1976. By E.
Patschelet. 2nd edition. Springer-Verlag, New York.
643 pp. Paper US $11.95; cloth US $27.60.
+Migratory game bird hunters and hunting in Canada. 1978.
Edited by H. Boyd and G. H. Finney. Canadian Wildlife
Service Report Series Number 43. Supply and Services
Canada, Ottawa. 125 pp., illus. $7.50 in Canada; $9
elsewhere.
The naturalists: pioneers of natural history. 1979. By Alan
K. Jenkins. Nelson, Toronto. 185 pp., illus. $17.95.
BOOK REVIEWS
353
+The outer shores: part 1, Ed Richetts and John Steinbeck
explore the Pacific coast. 1978. Edited by Joel W. Hedg-
peth. Mad River Press, Eureka, California. xii + 128 pp.,
illus.
+Pest management. 1978. Edited by G. A. Nortonand C. S.
Holling. Proceedings of a Conference, October 1976,
Austria. Pergamon, Oxford. 358 pp., illus. US $40.
Phosphorus in the environment: its chemistry and bio-
chemistry. 1978. By numerous authors. Ciba Foundation
Symposium 57 (New Series). Elsevier, Amsterdam. c300 pp.,
illus. Dfl. 72.60.
Resources and planning. 1979. Edited by B. Goodalland A.
Kirby. Pergamon, Elmsford, New York. 373 pp., illus. Cloth
US $33; paper US $15.
Sun: mankind’s future source of energy. 1979. Edited by F.
deWinter. Pergamon, Elmsford, New York. 3 volumes,
2250 pp. US $250.
The tides of the planet Earth. 1978. By Paul Melchior.
Pergamon, New York. 609 pp. US $75.
Yukon-Kuskokwim Delta. 1979. By various authors.
Alaska Geographic Volume 6, Number |. Alaska Northwest,
Anchorage. 96 pp., illus. US $11.95.
Wood as an energy resource. 1978. By D. A. Tillman.
Academic Press, New York. xiv + 252 pp., illus. US $13.50.
*Assigned for review
+Available for review
Call for nominations for the Council of The Ottawa Field-Naturalists’ Club
A Nominating Committee has been chosen by the
Council to nominate persons for election to offices
and membership of the Council for the year 1980, as
required by the Constitution.
Club members may also nominate candidates as
officers and other members of Council. Such nomi-
nations require the signatures of the nominator and
seconder, and a statement of willingness to serve in the
specified position by the Nominee. Nominations
should be sent to the Nominating Committee, The
Ottawa Field-Naturalists’ Club, Post Office Box 3264,
Postal Station C, Ottawa, Ontario K1Y 4J5, to arrive
no later than 15 November 1979.
The Committee will also consider any suggestions
for nominees which members wish to submit to it by |
November 1979. It would be helpful if some relevant
background on the proposed nominees were provided
along with the suggested names.
Chairman, Nominating Committee
Instructions to Contributors
Content
The Canadian Field-Naturalist is a medium for the
publication of scientific papers by amateur and professional
naturalists or field-biologists reporting observations and
results of investigations in any field of natural history
provided that they are original, significant, and relevant to
Canada. All readers and other potential contributors are
invited to submit for consideration their manuscripts
meeting these criteria. As the journal has a flexible
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Please submit, in either English or French, three complete
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Type the manuscript on standard-size paper, if possible
use paper with numbered lines, double-space throughout,
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Articles also require a running head. Generally words should
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Underline only words meant to appear in italics. The names
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Although we prefer the names of journals in the Literature
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Reviewing Policy of The
Manuscripts submitted to The Canadian Field-Naturalist
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The Council of Biology Editors Style Manual, 4th edition
(1978) available from the American Institute of Biological
Sciences, is recommended as a guide to contributors.
Webster’s New International Dictionary and le Grand
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Canadian Field-Naturalist
comments and constructive recommendations. Almost all
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TABLE OF CONTENTS (concluded)
Eastern limit of the Five-lined Skink, Eumeces fasciatus, in Ontario
RICHARD D. USSHER and FRANCIS R. COOK
Barn Owls in Quebec NORMAND DAVID
Communal roosting of Song Sparrows under snowbank MARTIN K. MCNICHOLL
Status of Eastern White Cedar, Thuja occidentalis, in western Nova Scotia GORDON S. RINGIUS
Snowy Egret in the Northwest Territories RICHARD W. QUINLAN
Letters
Tribute to Hoyes Lloyd, 1888-1978 DAvID A. MUNRO
News and Comment
Book Reviews
Zoology: Animals in peril: a guide to the endangered species of Canada and the United States — The
great arc of the wild sheep — A guide to the birds of Venezuela — North American Moose —
Wild geese — Crows of the world — Life of the Harp Seal.
Botany: Edible garden weeds of Canada — Ferns of the Ottawa district — Dyes from lichens and
plants: a Canadian dyer’s guide.
Environment: Human activity and the Environment — Vermillion Lakes Banff National Park: an
introductory study.
Miscellaneous: Weather almanac — Introduction to mathematics for life scientists
New Titles
Mailing date of previous issue 23 April 1979
1979 Council — The Ottawa Field-Naturalists’ Club
President: Roger Taylor Elisabeth Beaubien Fran Goodspeed
hah Charlie Beddoe Peter Hall
Vice-President: Courtney Gilliatt Ron Bedford Hue MacKenzie
Treasurer: Barry Henson Frank Bell Frank Pope
Bill Cody lan Sneddon
Recording Secretary: Diana Laubitz Jane Diceman Ken Strang
: : Ellaine Dickson Ken Taylor
C ov ;
orresponding Secretary: Valerie Hume Honey icieon eee ee
Past President: Roger Foxall Marc Forget
321
323
325
326
329
330
331
333) 7/
359
344
347
348
350
Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists’ Club,
Box 3264, Postal Station C, Ottawa, Canada KIY 4J5. For information on Club activities telephone (613) 722-3050.
THE CANADIAN FIELD-NATURALIST Volume 93, Number 3 1979
Editorial
To a Bigot LORRAINE C. SMITH
Articles
Seasonal growth, food and feeding habits of young-of-the-year Black
Crappie in the Ottawa River JOHN MARK HANSON and S. U. QADRI
Nesting and food-storage behavior of Peromyscus maniculatus gracilis and
P. leucopus noveboracensis C. CRAIG TADLOCK and HAROLD G. KLEIN
Life history characteristics of Little Brown Bats (Myotis lucifugus) in Alberta
DAVID B. SCHOWALTER, JOHN R. GUNSON, and LAWRENCE D. HARDER
Winter denning of the Striped Skunk in Alberta
JOHN R. GUNSON and RONALD R. BJORGE
Range extensions of vascular plants in northern Yukon Territory and
northwestern District of Mackenzie
JOHN A. NAGY, ARTHUR M. PEARSON, BERNARD C. GOSKI, and WILLIAM J. CODY
Food and feeding of the Rainbow Smelt (Osmerus mordax) in Lake Simcoe,
Ontario HUGH R. MACCRIMMON and ROBERT W. PUGSLEY
Song pattern of the Cypress Hills population of White-crowned Sparrows
M. Ross LEIN
Nesting biology and development of young in Ontario Black Terns ERICA H. DUNN
Summer, autumn, and winter diets of Elk in Saskatchewan - HUGH M. HUNT
Taxonomy, distribution, and ecology of the cliff-brake ferns ( Pellaea:
Polypodiaceae) in Alberta DANIEL F. BRUNTON
Characteristics of Peregrine Falcons migrating through central Alberta,
1969-1978 DIcK DEKKER
Notes
Extralimital record of a Narwhal (Monodon monoceros) in Hall’s Bay, Newfoundland
BorRA MERDSoyY, JON LIEN, and ANNE STOREY
First Canadian record of the Black Buffalo (Osteichthyes: Catostomidae)
E. J. CROSSMAN and S. J. NEPSZY
Thaspium trifoliatum (Meadow-parsnip) in Canada P. W. BALL
Largest Gray Wolf skulls found in Alberta JOHN R. GUNSON and RONALD M. NOWAK
Aquatic feeding by a Woodchuck D. FRASER
Dispersion of freshwater leeches (Hirudinoidea) to Anticosti Island, Quebec
RONALD W. DAVIES
Relative efficiencies of Museum Special, Victor, and Holdfast traps for sampling
small mammal populations ARTHUR M. MARTELL
Probable hybrids of Cinnamon X Blue-winged Teal from southern Alberta
D. V. WESELOH and LINDA MCKEANE WESELOH
Common Garter Snake predation on Ring-billed Gull chicks PETER M. FETTEROLF
The Rock Vole (Microtus chrotorrhinus) as a Transition Zone species
GORDON L. KIRKLAND, Jr. and CHARLES M. KNIPE
Zl
232
316
ST)
319
concluded on inside back cover
ISSN 0008-3550
aay
The CANADIAN ges
HARV
ote A i
UNIVERSI Ty
FIELD-NATURALIST
Published by THE OT
TAWA FIELD-NAT
- URALISTS’ CLUB
, Ottawa, Canada
The CANADIAN
FIELD-NATURALIST
ublished by THE INA Sis 2 5 awa, a
D-NATI STS
-D-NA RALI CLUB
Pi THE OTTAWA FIEI Ottawa, Canada
Contents
" Hibernation of bats in southeastern Ontario and adjacent Quebec.
By Harold B. Hitchcock 47
“Christmas bird eas 8 rn 59
. Propo als in potanical nomenclature. By Bernard Bova 66
Apparent observations of the whooping crane gn Central Saskatchewan.
By Lawrence Ho Walkinshaw as 73
A new Antennaria from Sorthern Ungava- By A. E. Fors d 80
"Salix petioloris 4. Smith in Edmonton Pisirict of Alberts.
By George ovum Be $2
The accurrence oi the leech Batrachabdella picta (Verrill) in the dorsal sub-cutane-
‘ous lymph spaces of Rana catesbiana. By Laurence R. Richardson - 85
‘Squirrel cache of funei By George A. mady se
Present state of the chestnut, Castanea dentata (Marsh.) Borkh., 1» Ontario.
C By W. Sherwood De Oe 3s
A puzzling Labrador reference. By W. L. MeAtee a9
Notes and Observations: — :
| . “Breeding of the Sora in New Bumswick. By George F. Boyer de 90
. Ground-nesting robins. By W- H. Whitehead” oe pe OL
Rana p. pipiens. By T. 9. Norris-Elye. < 91
‘Breeding killdeers in Northern New Brunswick.
‘ By Austin Ww. Cameron. 92
Book Reviews Oe ee 92
Published by the
7 OTTAWA FIELD-NATURALISTS CLUB
Entered at ‘the Post Office at Olawe, Ont., as second class matter, ‘ Volume 91, Number 1
January-March 1977
Centennial Year
Volume 93, Number 4
October-December 1979
The Ottawa Field-Naturalists’ Club
FOUNDED IN 1879
Patrons
Their Excellencies the Governor General and Mrs. Edward Schreyer
The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural
heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse
information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving,
maintaining or restoring environments of high quality for living things.
The Members of Council are listed on the inside back cover.
The Canadian Field-Naturalist
The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas
expressed in this journal, however, are private and do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or
any other agency.
Editor: Lorraine C. Smith
Assistant to the Editor: Donald A. Smith Book Review Editor: J. Wilson Eedy
Associate Editors
Gy Dy Bird A. J. Erskine George H. La Roi
E. L. Bousfield Charles Jonkel David P. Scott
Francis R. Cook Charles J. Krebs Stephen M. Smith
W. O. Pruitt, Jr.
Copy Editor: Marilyn D. Dadswell Chairman, Publications Committee: J. K. Strang
Production Manager: Pauline A. Smith Business Manager: W. J. Cody
Subscriptions and Membership
Subscription rates for individuals are $10 per calendar year. Libraries and other institutions may subscribe at the rate
of $20 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $10 includes a subscription to The
Canadian Field- Naturalist. Subscriptions, applications for membership, notices of changes of address, and undeliverable
copies should be mailed to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada KIY 4J5.
Second Class Mail Registration No. 0527 — Return Postage Guaranteed.
Back Numbers
Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field- Naturalists’ Club, 1879-
1886, and The Ottawa Naturalist, 1887-1919, may be purchased from the Business Manager.
Business Manager: Mr. W. J. Cody, Box 3264, Postal Station C, Ottawa, Ontario, Canada KIY 4J5
Book Review Editor: Dr. J. Wilson Eedy, R.R. 1, Moffat, Ontario LOP 1J0
Coordinator, The Biological Flora of Canada: Dr. George H. La Roi, Department of Botany, University of Alberta,
Edmonton, Alberta T6G 2E9
Address manuscripts on birds to the Associate Editor for Ornithology:
Dr. A. J. Erskine, Canadian Wildlife Service, Box 1590, Sackville, New Brunswick EQOA 3C0
All other material intended for publication should be addressed to the Editor:
Dr. Lorraine C. Smith, R. R. 3, Stittsville, Ontario, Canada KOA 3G0
Urgent telephone calls may be made to the Editor's office (613-996-5840), the office of the Assistant to the Editor (613-231-
4304), or their home on evenings and weekends (613-836-1460), or to the Business Manager's office (613-995-9461).
Cover: Since The Ottawa Field-Naturalists’ Club was founded in 1879, it has published a scientific journal. Covers of earlier
issues of The Canadian Field- Naturalist are illustrated: the contents were listed on covers from 1945-1969: the larger
format with a photograph was introduced in 1970.
The Canadian Field-Naturalist
Volume 93, Number 4
October-December 1979
Some New and Interesting Vascular Plant Records from
Northern Ontario
J. L. RILEY
Department of Botany, Royal Ontario Museum, Toronto, Ontario
MS5S 2C6
Riley, J. L. 1979. Some new and interesting vascular plant records from northern Ontario. Canadian Field-Naturalist
93(4): 355-362.
Locations and habitats are described for recent additions to the native arctic-subarctic flora of Ontario, and for selected
subarctic species which had been previously known from only one or two sites. Additions to the provincial flora are Agrostis
borealis, Aster alpinus, Calamagrostis deschampsiodes, Carex bigelowii, Hierochloe alpina, Kobresia myosuroides, Luzula
confusa, Minuartia groenlandica, Phyllodoce coerulea, and Vahlodea atropurpurea. Comments are offered on the occur-
rence of more widespread species such as Hierochloe pauciflora, Kobresia simpliciuscula, Pinguicula villosa, Agropyron
violaceum, Ledum decumbens, and arctic-subarctic species of the genus Pedicularis. Cypripedium calceolus var. plani-
petalum is also recorded.
Key Words: northern Ontario, vascular plants, arctic-subarctic, phytogeography, mapping, rare or restricted.
During fieldwork directed towards a compilation of
the vascular flora of the Hudson Bay Lowland, over
350 different Ontario sites have been visited. Many
species new to this flora have been documented. The
most interesting group are those restricted to the
Hudson Bay area. The purpose of this paper is to
report and discuss these new records and the
distributions of restricted species now recorded from
significantly more sites, in order to provide a useful
data base from which to speculate on the character
and extent of Ontario’s “maritime tundra.”
Species Accounts
Agrostis borealis
Carex bigelowii
Hierochloe alpina
Luzula confusa
Minuartia groenlandica
Kenora District: 54°28’N, 84°54’W:; summit of ab-
ruptly rising Precambrian instrusive cuesta west
of Hawley Lake, Birch Hill. Riley, 29 July 1978.
54°18’N, 84°30’W: summit of ab-
ruptly rising Precambrian intrusive cuesta east
of Aquatuk Lake, elevation 900+ ft. [275 m] asl
Riley, 29 July 1978.
The Sutton Ridges are a discontinuous series of
Precambrian outcrops of various elevations and
exposures extending across the northeastern Hudson
Bay Lowland from the Nowashe River watershed
northwestward to within 20 km of the Hudson Bay
coast just east of Winisk. These prominent outcrops
(Figure 1) are Precambrian diabase-gabbro sills
overlying sedimentary Precambrian rocks consisting
of a lower dolomitic unit overlain by a unit consisting
of iron formations, greywacke, and other materials
(Bostock 1968). Two of the most prominent of these,
Birch Hill and Aquatuk Lake, had open rock barren
summits supporting arctic species characteristic of
such Precambrian habitats in northern Quebec. Other
outcrops, such as at the Sutton-Hawley Lakes
Narrows and near the Hudson Bay coast (55°N,
84°43’W) apparently do not support such species,
although the flora of the former includes arctic-
subarctic species such as Potentilla nivea (known
elsewhere in Ontario only from near the lower.
Shagamu River: Sims 2258; SSM, TRT) and from
Cape Henrietta Maria (CAN) (Figure 1).
As well as providing suitably exposed upland
outcrop habitat, these summits were the first islands
to emerge from the postglacial Tyrrell Sea in the
northeastern Lowland, about 6000 yr BP (Webber et
al. 1970). As a result, they have been available to the
long-distance dispersal of plants for a much longer
period of time than any of the surrounding Lowland
except for areas far to the south and west, where none
of these species now persist.
Agrostis borealis (Riley 9511, 9464) is a circum-
polar low arctic species characteristic of “dry rocky
and turfy places on acid rock” (Porsild 1957) and of
“late-snow-free areas” (Hultén 1968). It was mapped
by Porsild (1964) from the northeastern Hudson Bay
Lowland but without apparent voucher reference
355)
356 THE CANADIAN FIELD-NATURALIST
Vol. 93
FIGURE |. The Sutton Ridges, a series of Precambrian outcrops penetrating the Paleozoic wetlands characteristic of the
Hudson Bay Lowland. This Aquatuk Lake ‘cuesta’ has an elevation ca. 275 m asl.
(CAN, DAO, TRT) nor with any mention made of it
in his northern Ontario fieldnotes (CAN). It is not
included by Scoggan (1978) in the flora of Ontario.
The specimens collected (CAN, TRT) are typically
awned, with the narrow leaves of var. borealis. It is
also known from a third Ontario location (55°N,
84°43’W: abandoned Mid-Canada Line radar station
510, occupying a small inlier of Precambrian diabase-
gabbro rock; Sims 2462A, 10 August 1978 (SSMF,
Xerox TRT)) (Figure 2).
Carex bigelowii ( Riley 9536, 9460) (Figure 2) is an
amphi-Atlantic member of an arctic circumpolar
Species complex (refer map, Hultén 1964), charac-
teristic of dry tundra and rocky habitats. Although
also mapped by Porsild (1964) from the northeastern
Hudson Bay Lowland, Scoggan (1978) reports it
absent from the Ontario flora. Again, no reference is
made to it in Porsild’s fieldnotes from northern
Ontario (CAN), nor is it represented in major Ontario
herbaria (CAN, DAO, TRT). The Birch Hill (CAN,
TRT) and Aquatuk Lake (TRT) specimens are
typical; collections from outcrop near the Hudson
Bay coast (55°04’N, 84°50’W; Riley 9666, 31 July
oe ae
Luzula confusa
Carex bigelowii
ee Agrostis borealis
_ ,» Minuartia groenlandica
Pay
Ty
& eA 5 r .
0 2 4 6 80 100 M1
SS
o @ 8 I~ wo Km
‘ (
te
a *.
yl
at
FIGURE 2. Ontario distribution of Luzula confusa, Carex
bigelowii, Agrostis borealis, and Minuartia groen-
landica.
1979
1978 (TRT)) and from willow thickets near North
Point, James Bay (51°28’N, 80°27’W; Tessier, 15 July
1976 (TRT), det. Ball) confirm Porsild’s (1957)
observation that the species’ habit is “more lax in
wetter sites” (Figure 2).
Hierochloe alpina (Riley 9458) was collected only
on the summit of the outcrop at Aquatuk Lake(CAN,
TRT) (Figure 3). Specimens collected belong to the
subspecies orthantha, having untwisted awns attached
above the middle of the second glume (Weimarck
1971). This taxon occurs from Greenland to the east
coast of Hudson Bay and James Bay south to several
mountain areas of New England and upper New York
State. It reoccurs sporadically in Beringia, but this
collection represents the western limit of its major
range; the closest westward location, at Cape
Churchill, is of ssp. a/pina. This collection represents
the first Ontario record of this circumpolar arctic-
alpine species. John Macoun’s reference (1888) to its
occurrence at Moose Factory, James Bay, is un-
substantiated and highly improbable (Dutilly et al.
1954) (Figure 3).
Luzula confusa (Riley 9512, 9467) is a circum-
polar arctic species, occurring southward in the
Canadian Rockies and ona few mountain tops in New
England, in dry turfy tundra heath, and on rocky
slopes and ledges (Porsild 1957; Hultén 1968). These
first Ontario records (TRT) complete the distribu-
tional gap between occurrences at Churchill, Mani-
toba and the northeastern shore of James Bay,
Quebec (Figure 2).
Minuartia groenlandica (Arenaria g.) ( Riley 9526,
9471), the attractive Greenland sandwort, is an
82°
80 100 MI
120 160 Km
e@ Hierochloe alpina
a ~ Agropyron violaceum
% Calamagrostis deschampsioides
%t Aster alpinus
FIGURE 3. Ontario distribution of MHierochloe alpina,
Agropyron violaceum, Calamagrostis deschamp-
sioides, and Aster alpinus.
RILEY: NEW NORTHERN ONTARIO PLANTS
357
eastern North American subarctic endemic, occurring
from Greenland, southwestward through Newfound-
land, Nova Scotia and in New England and northern
New York State mountains, northwestward to the
east coast of Hudson Bay north of James Bay, and
now known as far westward as the Sutton Ridges in
Ontario (for Quebec distribution, refer to Rousseau
1974). As with the other arctic-subarctic species
disjunct on outcrop summits of the Sutton Ridges, the
nearest major ranges of species are across the mouth
of James Bay, in the Precambrian uplands of northern
Quebec (Figure 2).
Agropyron violaceum (Hornem.) Lange (A. /atiglume
(Scribn. & Sm.) Rydb.)
Kenora District: Vicinity of the mouth of the Severn
River. Moir 1416, 14 Aug. 1952 (CAN) (sub A.
trachycaulum var. novae-angliae; rev. Porsild
1959).
Vicinity of the mouth of Goose Creek. Moir,
18-20 Aug. 1952 (CAN) (sub A. t¢. var. n.-a.: rev.
Porsild).
15 mi [25 km] from the mouth of the Black
Duck river. Moir 2087, 25-27 July 1953 (CAN)
(sub A. t. var. n.-a.; rev. Porsild 1959).
56°00’N, 87°38’W; Fort Severn, near H.B.C.
post. Hustich 1256, 28 July 1956 (CAN) (det.
Porsild).
55°14’N, 85°05’W; Winisk, disturbed gravel-
clay areas near airport. Riley 5872, 24 August
1976 (TRT).
55°14’N, 85°07’W, Winisk airport area, near
fuel depot. Riley 6/40, 26 August 1976 (TRT).
55°52’N, 86°46’W; coast of Hudson Bay, im-
mediately east of mouth of Shagamu River. Riley
7483, 11 August 1977 (TRT).
55°57’'N, 89°57’W; littoral habitat along
Black Duck River. Riley 8693b, 19 July 1978
(TRT).
55°N, 84°43’W: abandoned Mid-Canada
Line radar station 510. Sims 2472, 10 August
1978 (SSMF, TRT).
55°04’50”N, 85°39’30”W; upland thicket along
river levee. Sims 2/30, 23 July 1978 (SSMF,
IRA).
This species, first reported for Ontario by Hustich
(1957. Comparison of the flora and vegetation in the
Fort Severn and in the Big Trout area. 2nd report
concerning fieldwork in 1956. Arctic Institute of
North America, University Library Tower, 2920-
24th Ave., N.W., Calgary, Alberta), occurs north to
Ellesmere Island and is transcontinental from Alaska
to Greenland, south to northern Manitoba and
Ungava; these specimens represent southern limits for
this arctic species (refer map, Porsild 1964). None of
the specimens seen have the pubescent glumes of the
358
more northern variety hyperarcticum. At Winisk it
occurs in disturbed gravel areas, having colonized
them since the air base was built in the 1950s. The
other locations are from riverbanks of major spring-
flooding rivers, in habitats such as low shrub willow
thickets (Riley 7483; Hustich 1957) and graminoid
meadows. Such calcareous alluvial sites are among
the most floristically rich in the coastal area of
southwestern Hudson Bay; the author has recorded
up to 60 species within 30 m of closed and open low
willow thickets along the lower Shagamu River
(Figure 3).
Aster alpinus
Kenora District: 55°52’N, 86°46’W: coast of Hudson
Bay, mouth of Shagamu River. Riley 6972, 29
July 1977 (CAN, TRT).
This improbable find was made among the gravel
bars at the mouth of the Shagamu River, where it grew
with such species as Chrysanthemum bipinnatum ssp.
huronense, Epilobium latifolium, and Elymus mollis.
It represents the first record of the species between the
Great Slave Lake area and the Pyrenees in Europe.
The North American phase, ssp. vierhapperi, is widely
distributed in the Rocky Mountains through to
Siberia (refer map, Hultén 1968) (Figure 3).
Calamagrostis deschampsioides
Kenora District; 55°44’10”N, 86°19’55”W; SW
coast of Hudson Bay, about 15 km SW of the
Shagamu River mouth. Sims 2245, 28 July 1978
(SSMF, TRT).
This Ontario record for this subarctic circumpolar
coastline species helps to confirm its distribution
around Hudson Bay, filling a gap between Churchill
and York Factory, Manitoba (Scoggan 1957), and
northern James Bay and Hudson Bay, Quebec
(Rousseau 1974; Hultén 1968). It was found in a
brackish marsh dominated by Carex aquatilis and
Scirpus hudsonianus, about 0.5 km above the high-
tide level, in peat 38 cm deep (Figure 3).
Cypripedium calceolus var. planipetalum
Kenora District: 52°57’30’N, 83°10’W; limestone
outcropping across the Attawapiskat River
about 40 mi [65 km] upstream of Attawapiskat
townsite. Riley 6328, 3 July 1977 (TRT).
52°52’N, 83°45’W: northward facing lime-
stone outcrop ... along south shore Attawa-
piskat River. Riley 6573b, 5 July 1977 (TRT).
This variety was first described by Fernald (1926)
from “turfy limestone barrens,” “peaty . . . lime-
stone barrens,” etc., in northwestern Newfoundland.
It was distinguished by its broad and short sepal,
broad flattish petals, and a more cordate stamino-
dium. Luer (1975) recognizes the taxon, although his
distribution map inexplicably extends its range from
THE CANADIAN FIELD-NATURALIST
Vol. 93
Newfoundland up the St. Lawrence River to eastern
Ontario.
It was found growing in open limestone White
Spruce (Picea glauca) forest along the lower Attawa-
piskat River, associated with Juniperus horizontalis,
J. communis, Alnus crispa, and Arctostaphylos uva-
ursi. In the second site, it was found close to var.
pubescens (Willd.) Corr. It is suggested that the status
of this taxon requires closer consideration, especially
in sites in which it is sympatric with other intraspecific
taxa.
Hierochloe pauciflora
Kenora District: 54°40’N; 34 mi [55 km] au sud du
cap Henrietta Maria. Dutilly, Lepage 3141/1,
17-18 August 1953. (Dutilly et al. 1954).
55°07’N; Cap Henrietta Maria. Dutilly,
Lepage 31124, 10 August 1953. (Dutilly et al.
1954).
55°52’N, 86°48’W; mouth of Shagamu River,
two miles inland, one mile west of river. Riley
294, 3 August 1977 (TRT).
56°47’N, 88°53’W: West Pen Island. Riley
8438, 16 July 1978 (TRT).
55°N, 82°18’W; Cape Henrietta Maria. Riley
9292, 27 July 1978 (TRT).
55° 14’N, 84°49’10”W; SW coast of Hudson
Bay, approx. 10 km E of Winisk, 1.25 km from
coast. Sims 2417, 9 August 1978 (SSMF, Xerox
TRIP).
This high arctic species (Porsild 1957), circumpolar
except for Greenland and Scandinavia, is a wet tundra
associate. Porsild’s observation (1957), that the
species is, “although not littoral, a distinctly coastal
and lowland species,” is apparently true along
Ontario’s Hudson Bay coast. It is found in mossy
inter-ridge tundra situations, such as at Cape Hen-
rietta Maria, and in inter-ridge mossy willow thickets.
It appears to achieve highest dominance values in
open graminoid fen-meadows close to the coast with
high pH values (6.2 and 7.2 recorded) and shallow
peats (30 to 45cm over permafrost or silts). One
collection (Riley 8438) was made in a graminoid
association dominated by Ca/amagrostis stricta, with
Hierochloe pauciflora having cover value approach-
ing 10%. Other species present at this site included
Carex rariflora, Pedicularis sudetica, Salix candida,
S. brachycarpa, and Saxifraga hirculus (Figure 4).
Kobresia myosuroides (Vill.) Fiori & Paol. (K.
bellardii (All.) Degl.)
Kenora District: 55°52’N, 86°46’W; coast of Hudson
Bay, immediately east of mouth of Shagamu
River. Riley 74806, 11 August 1977 (TRT).
55° 22’35”N, 85° 22’50” W; SW coast of Hud-
son Bay approx. 7 km NW of Winisk, 2 km from
84° 82
Hierochloe pauciflora
coast. Sims 2180b (mixed coll’n with Carex
bicolor All.), 26 July 1978 (TRT).
Easily overlooked, this tiny sedge is undoubtedly
more widespread along Ontario’s Hudson Bay coast.
A circumpolar arctic-subalpine species occurring to
northernmost Ellesmere Island, it is “a pronounced
calciphile, preferring dry, sandy heath and windswept
ridges” (Porsild 1957). The Shagamu River collection
was from a silted, riparian low willow thicket
association, while the Winisk collection was from an
immature low shrub thicket with shallow peat (7 cm) ,
and a dense graminoid ground cover including Carex
microglochin, C. rariflora, C. capillaris, Scirpus
cespitosa, and Kobresia simpliciuscula (Figure 5).
2 0 20 © 6 80 100 Mi
ee oe
o 0 44 80 120 160 Km
4 Kobresia myosuroides
e K. simpliciuscula
FIGURE 5. Ontario distribution of Kobresia myosuroides
and K. simpliciuscula.
RILEY: NEW NORTHERN ONTARIO PLANTS
358
Kobresia simpliciuscula
Kenora District: 55°07’N; Cap Henrietta Maria.
Dutilly, Lepage 31135, 10 August 1953 (Dutilly
et al. 1954).
54°34’N, 84°38’W; Hawley Lake, Sjors,
1961 (Lepage 1966).
54°48’N, 82°23’W; Runway 415, south of
Cape Henrietta Maria, Riley 5850, 23 August
1976 (TRT).
56°22’N, 88°08’W. Riley 8465, 16 July 1978
(TRT).
55°29’10”N, 85°41’1S5”W: SW coast of
Hudson Bay, approx. 16km NW of Winisk,
7.25 km from coast. Sims 2043, 20 July 1978
(SSMF, TRT).
55° 22’35”N, 85°22’50”W; SW coast of Hud-
son Bay, approx. 7 km NW of Winisk, 2 km from
coast. Sims 2182, 2185, 26 July 1978 (SSMF,
(TRT).
55°02’N, 82°41’W. Riley 9169, 27 July 1978
(TRT).
54°47’'N, 82°39'W. Riley 9358, 28 July 1978
(TRT).
54° 50’N, 82°46’W. Riley 9372, 28 July 1978
(TRT).
55° 12’N, 84° 17’'W. Riley 9401, 29 July 1978
(TRT).
54° 19’N, 84°30’W. Riley 9476, 29 July 1978
(TRT).
54° 38’N, 84°37’W; slope of Sutton River
about | km from the river. Riley 9567, 29 July
1978 (TRT).
54° 38’N, 85° 20’W. Riley 9631, 30 June 1978
@iRsh):
53° 38/Ni 83229. W. Riley, 9737, 3 August
1978 (TRT).
The distribution in Ontario (Figure 5) of this
circumpolar arctic-alpine sedge is much more wide-
spread than indicated by the few early records. The
dozen subsequent records are all from calcareous
habitats; six were made from open graminoid or open
low shrub fens with pH’s between 5.8 and 7.2, peat
depths from 7 to 50cm. These fen sites all had a
vascular plant ground cover dominated by Scirpus
cespitosa, with Carex rariflora, C. livida, C. limosa,
Betula glandulifera, Juniperus horizontalis, and
Myrica gale as common associates. The most com-
mon bryophytes recorded were Scorpidium scor-
poides, Tomenthypnum nitens, and Drepanocladus
spp. The non-wetland sites in which Kobresia sim-
pliciuscula occurs include a variety of ericaceous
tundra heath, frost-shattered limestone, frost-sorted
till, and algal marl habitats not uncommon through
the interior of the Hudson Bay Lowland in an area
extending from the lower Ekwan River to the lower
Winisk, northeastward to Cape Henrietta Maria.
360 THE CANADIAN FIELD-NATURALIST
Current studies in such interior areas confirm that
terrestrial arctic habitats are not confined simply to a
narrow coastal strip but extend to non-arboreal
terrestrial (usually till upland or bedrock outcrop)
habitats over a much wider area.
Phyllodoce coerulea
Kenora District: 55°07’N, 83°52’35”W; SW coast of
Hudson Bay approx. 6 km SW of Sutton River
mouth, 16 km from coast. Sims 2397a, 4 August
1978 (SSMF, TRT).
The closest records of this circumpolar subarctic-
alpine mountain heather are from Cape Jones,
Quebec (Rousseau 1974) to the east and from the
Precambrian Shield of northern Manitoba to the west
(Scoggan 1957; Ritchie 1962). It was collected from an
upland open White Spruce-lichen woodland, with
trees reaching 15 m (230 yr old), shallow lichen peat
(less than 10 cm), and a water table ca. 50 cm deep.
Lichen woodland on such beachridge complexes
appears to burn regularly; those supporting forests
older than 200 yr should be considered very rare
(Figure 6).
Cody (1953) mapped and discussed its North
American distribution and concluded that P. coerulea
has widely re-occupied its glaciated northeastern
range. Its occurrence 16 km from the Hudson Bay
coast suggests a maximum age for the migration of the
species into Ontario. The site is estimated to have
undergone postglacial isostatic uplift over the last
1000 yr of about 1.2 m per century, and to have been
above sea level for only about 1500 yr (Webber et al.
1970).
84°
5; : ae
Z Phyllodoce coerula
A | “( 0 Ledum decumbens
Vol. 93
Pinguicula villosa
Kenora District: 54°34’N, 84°38’W; Hawley Lake,
Sjors, 1961 (Lepage 1966).
56° 16’N, 88°05’W. Riley 8478, 16 July 1978
(TRT).
56° 48’N, 88°52’W. Riley 8485, 16 July 1978
(ERD):
55°48’N, 87°31’W. Riley 9042, 23 July 1978
(TRT).
55°20'05’N, 85°39’15”W; approx. 12 km W
of Winisk, 17 km from coast. Sims 2089, 13 July
1978 (SSMF, TRT) (sub. P. vulgaris).
55° 11’N, 85°40’W. Riley 9/42, 26 July 1978
(TRT).
54° 28’N, 84°54’W. Birch Hill. Riley 95/3, 29
July 1978 (TRT).
54° 38’N, 85°20’W. Riley 9633, 30 July 1978
(TRT).
This circumpolar subarctic butterwort is not an
infrequent acidophile of the fen and bog wetlands
developing well back of the Hudson Bay coast (Figure
7). Within open graminoid fen patterns of the
lowlands, it is found on bog (sphagnum) hummocks,
palsas or on the bog ‘laggs’ or ‘bibs’ of peat plateau
patterns. On the summit of Birch Hill, it was found
growing beside a confined wet sphagnum pool.
f 7h a
2 0 2 40 60 80 100 ml |
SSeS eS ee
0 4 80 12 160 Km |
we ie
fi 3 |
Vahlodea atropurpurea
Kenora District: 55°07’N, 83°52’35”W; SW coast of
Hudson Bay approximately 6 km Sw of Sutton
River mouth, 16 km from coast. Sims 2397, 4
August 1978 (SSMF, Xerox TRT) (det. Rez-
nicek).
This subarctic circumpolar grass is known as a rare
tundra species in Manitoba (Scoggan 1957, 1978) and
0 2 4 6 80 00 ml ~”
4 80 120 wo Km
Ay F
FIGURE 6. Ontario distribution of Phyllodoce coerula and
Ledum decumbens.
IT)
a widespread species of Quebec’s alpine meadows and
open conifer forests on sandy soils (Rousseau 1974).
This first Ontario record is from an open White
Spruce-lichen woodland site also supporting Phyl-
lodoce coerulea (see further site comments with that
species) (Figure 6). This specimen is of the typical,
eastern variety.
Recent field work in the Hudson Bay Lowland of
Ontario has thus resulted in the addition of 10
vascular plant species to the provincial flora. These
are Agrostis borealis, Carex bigelowii, Hierochloe
alpina, Luzula confusa, Minuartia groenlandica,
Vahlodea atropurpurea, Aster alpinus, Calamagrostis
deschampsioides, Kobresia myosuroides, and Phyl-
lodoce coerulea. The first five of these species are
reported from the highest and most exposed of the
outcrop summits of the Precambrian Sutton Ridges.
Other than their nearly complete inaccessibility, these
two stations do not currently enjoy any protective
status. The other species are all from widely dispersed
locations and, with the exception of Cypripedium
calceolus var. planipetalum (lower Attawapiskat
River), all are now known from sites within the Polar
Bear Provincial Wilderness Park.
Discussion
All of the species reported here as new to the
provincial flora should be included in the listing of
rare vascular plants (Argus and White 1977). Of the
other taxa discussed or mapped, Pinguicula villosa,
Pedicularis labradorica, and Cypripedium calceolus
var. planipetalum should also be listed as rare.
Because of the rarity of suitable habitat in Ontario, it
appears that some species are undoubtedly rare:
Agrostis borealis, Carex bigelowii, Hierochloe alpina,
Luzula confusa, Minuartia groenlandica, Phyllodoce
coerulea, Aster alpinus, Cypripedium calceolus var.
planipetalum. Others are widespread but regionally
restricted: Pedicularis spp., Ledum decumbens,
Agropyron violaceum, Kobresia simpliciuscula, Pin-
guicula villosa. Some are species of greatly under-
sampled habitats: Hierochloe pauciflora, Calama-
grostis deschampsioides, Kobresia myosuroides.
In light of the currently accepted rough equivalence
between a ‘regionally restricted’ status and a ‘provin-
cially rare’ status, a caution may be in order. The
“narrow band of tundra along the Hudson Bay coast”
(Argus and White 1977) differs from the Carolinian
zone (Ontario’s southern zone of coincident restricted
species) in that its wildlife remains largely intact,
continuous and unendangered, and in that its geo-
graphic extent remains largely undefined, but almost
certainly far larger than the Carolinian zone of
southwestern Ontario. Mapping of most of Ontario’s
arctic-subarctic species confirms a ‘maritime tundra’
RILEY: NEW NORTHERN ONTARIO PLANTS
361
concept of Ontario’s Hudson Bay shore (Figure 4,
Hierochloe pauciflora; Figure 8, Pedicularis spp.).
The distributions of species such as Kobresia sim-
pliciuscula (Figure 5), Pinguicula villosa (Figure 7),
and Ledum decumbens (Figure 6), however, indicate a
much expanded zone.
2 4 60 80 100 mI
1eo Am
Sy
Pedicularis flammea
P. lapponica
P. labradorica
P. sudetica
2 ELS ar
FIGURE 8. Ontario distribution of Pedicularis flammea, P.
lapponica, P. labradorica, and P. sudetica.
Ritchie (1962) considered the subarctic zone in
Manitoba to be characterized by a vegetation of
primarily open conifer forest or a mixture of forest
and tundra, and by a climate with a growing season
not over 650 degree-days Centigrade and with a mean
July temperature not over 14°C. This included all of
the Hudson Bay Lowland in Manitoba, as well as the
Precambrian Shield westward. Meteorological data
from extreme northern Ontario are not presently of
sufficient detail to delimit such a climatic zone. The
‘openness’ of Hudson Bay Lowland associations is not
a function of dominant lichen woodland associations
or of tundra thinning forest sites, but the result of the
overwhelming dominance of wetland or peatland
vegetation on clay-silt marine substrates. Zonal
mapping by winter satellite photography (Hare and
Ritchie 1972) shows the Hudson Bay Lowland to have
albedo levels higher than, or comparable to wood-
lands, but this is not a result of any ‘openness’ of
terrestrial systems. For similar reasons, the standard
latitudinal zones, as summarized by Scoggan (1978),
are not valuable in proposing boundaries for an
arctic-subarctic floristic province in northern On-
tario. As the collected floristic data base is completed,
it is hoped that floristic criteria themselves will define
such a zone.
°362 THE CANADIAN FIELD-NATURALIST
Acknowledgments
This work was made possible through the field
support and encouragement of several individuals and
agencies; S. Pala and A.N. Boissonneau of the
Ontario Center for Remote Sensing; R. K. Ross and
R. I. G. Morrison of the Canadian Wildlife Service,
and D. W. Cowell of the Federal Lands Directorate,
Burlington. I especially thank R.A. Sims of the
Canadian Forestry Service, Sault Ste. Marie, for
sharing an enthusiasm for botanizing this wilderness
and for sharing his discoveries.
Mapping is based on specimens at CAN, DAO,
SSMF, and TRT (1978), and on records as indicated
in the text. J. M. Gillett (CAN) and W. J. Cody
(DAO) kindly checked herbarium specimens and the
field notes of A. E. Porsild (CAN) for references to the
species new to Ontario. P. W. Ball of Erindale
Campus, University of Toronto, assisted with the
identification of some sedge specimens, and J.
McNeill of the Biosysematics Research Institute,
Agriculture Canada, verified determinations of Agro-
pyron violaceum. Review of an earlier draft by R. A.
Sims and J. H. Soper was very much appreciated.
Literature Cited
Argus, G. W. and D. J. White. 1977. The rare vascular
plants of Ontario. National Museum of Natural Sciences,
National Museums of Canada, Syllogeus Series Number
14.
Bostock, H. H. 1968. Precambrian sedimentary rocks of
the Hudson Bay Lowlands. /m Earth science symposium
on Hudson Bay. Edited by P.J. Hood. Geological
Survey of Canada Paper 68-53: 206-214.
Cody, W. J. 1953. Phyllodoce coerulea in North America.
Canadian Field-Naturalist 67: 131-134.
Dutilly, A., E. Lepage,and M. Duman. 1954. Contribution
a la flore du versant occidental de la baie James,
Ontario. Contribution of the Arctic Institute, Catholic
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Received 22 January 1979
Accepted 23 April 1979
fags
Su ad
Yearly Variations in the Population Dynamics of
Richardson’s Ground Squirrels
GAIL R. MICHENER
Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E1
Present address: Department of Biological Sciences, University of Lethbridge, Lethbridge, Alberta T1K 3M4
Michener, G. R. Yearly variations in the population dynamics of Richardson’s Ground Squirrels. Canadian Field-Naturalist
93(4): 363-370.
Population fluctuations were followed from 1975 to 1978 in a group of Spermophilus richardsoniiin southern Alberta. Adult
males ranged widely in early spring but they restricted the extent of their activities once all females were mated, and
consequently fewer males were located on the study area after the breeding season than in early spring. Adult females made
only minor changes in the location of their ranges from year to year and the major loss of adult females occurred overwinter.
Each year fewer juveniles entered the active population than expected from the number of resident females and their
reproductive potential. There was net juvenile immigration in the year with the lowest initial density and net juvenile loss in the
years with highest density. Both predation and dispersal contributed to juvenile losses, particularly of males in 1976. The adult
sex ratio favored females regardless of the sex ratio of the juvenile population in the previous fall. Telemetric studies are
required to distinguish mortality in late summer from entry underground to hibernate and to distinguish death underground
during winter from early emergence in spring followed by dispersal or death.
Key Words: Ground squirrels, Spermophilus richardsonii, dispersal, overwinter mortality, sex ratio, population dynamics,
annual variations.
Large scale mark-release-recapture studies, by compare the results obtained by this method with
Dorrance (1974), Michener and Michener(1977),and those of previous studies.
Schmutz et al. (1979), of the population dynamics of
Richardson’s Ground Squirrels, Spermophilus Methods
richardsonti, have revealed that there is high loss of The Richardson’s Ground Squirrel population was
squirrels between years, that male losses are greater located on fescue grassland in Fire Guard Coulee
than female losses, that juvenile losses exceed adult (50°34’N, 114°18’W, elevation 1235 m)5.5 km NW of
losses, and that the adult sex ratio is biased infavor of Longview, Alberta. An area of 0.72 ha was gridded
females. Discrepancies exist between these workersin with flags at 20-m intervals, and a record was kept of
accounting for both the magnitude of losses and the the identity and numbers of squirrels, and of their
differential nature of losses. Dorrance (1974) con- location within this area. Squirrels were individually
cluded that overwinter mortality and loss of pups marked with a numbered metal tag in each ear and
prior to weaning were the main determinants of with black dye marks on the fur. As part of a
population size, whereas differential mortality favor- behavioral study conducted in 1975 all squirrels
ing adult females in early spring and favoring juvenile caught that year were further identified with a colored
females in fall and winter contributed to the biased _ plastic disc attached to one ear tag.
adult sex ratio. Michener and Michener (1977) Following initial trapping, which began on 13 April
concluded that overwinter mortality was a major 1975, the squirrels on the area were observed for 147 h
factor accounting for both the extent of interyear on 55d between 5 May and 11 October; in general,
losses and the differential nature of losses in their squirrels were retrapped only when redying of the
population. Schmutz et al. (1979) concluded that identifying marks was necessary. In 1976, squirrels
overwinter mortality, while reducing population size, | were observed for 32 h on 13 d between 28 Marchand
did not account for the disparate sex ratio but that 4 July. Thereafter, the area was trapped at approxi-
predation, differentially directed toward dispersing mately 2-wk intervals until 11 October to determine
adult males in spring and dispersing juvenile males in | which squirrels were still resident. In 1977 and 1978,
summer, did. all data were collected by trapping at approximately
The aim of this study was to examine the population _10-d intervals between 20 March and 2 October, and
dynamics of a small group of individually identifiable 19 March and 24 April, respectively. Under the
squirrels by observation and trapping at more observation regime the area was surveyed 5 times an
frequent intervals than used in previous studies inan hour for approximately 3 continuous hours, and all
attempt to distinguish residents from transients, to active squirrels were identified by their colored disc
follow the fate of each resident squirrel, and to and dye marks. Under the trapping regime an attempt
364 THE CANADIAN FIELD-NATURALIST
was made to capture every squirrel present; in 1976
this was done by placing an additional dye mark on
each captured squirrel until all squirrels were so
marked, and in 1977 and 1978 captured squirrels were
held in the live-traps until no other squirrels remained
active. As there were relatively few adult squirrels, all
of which could also be identified without capture,
errors in determining the number of residents were
likely only within the juvenile cohort. Juveniles were
sometimes not caught during one trapping episode
but captured during the next one; such animals were
assumed to have been in continuous residence and
were included in the population count for the day on
which they were not caught. On average, such ‘missed’
squirrels accounted for less than 8% of the juvenile
population present during any trapping episode.
Squirrels adjacent to the study area were trapped ona
sporadic basis each year.
Except where otherwise indicated, the term ‘resi-
dent’ refers to a squirrel present on the area for a
minimum of 2 wk, whose major burrow system was
within the boundaries of the area, and which spent the
majority of its-active time on any day within the area.
Vol. 93
Results
Adults
Adult males trapped in spring could be classed into
three categories: transients which were present for at
most a day or two before moving off the area,
temporary residents which were present on the area
for at least a week but did not remain resident for the
summer, and permanent residents which remained on
the area beyond the breeding season. Table | indicates
the yearly residency status of each male classed, in at
least one year, aS a permanent resident or as a
temporary resident. In the 4 yr of this study there was
only one instance of an adult male remaining as a
permanent resident in 2 consecutive years and one
instance of a male born on the area remaining resident
as a yearling. Three other yearlings that had been born
on the area were recovered; one was classed as a
transient in 1976 and two as temporary residents in
1977. Although at least half of the males classed as
permanent residents had moved on to the area in early
spring, there were no instances of adult male
immigration during the remainder of the active
season.
TABLE | —Yearly residency status of adult males classed as permanent or temporary residents in at least one year (see text for
criteria of classification). For those males also caught as juveniles their status as a juvenile is given
Residency status
Male 1975 1976 1977 1978
RO permanent
RB permanent
RBSS permanent
6201 temporary ;
6210 temporary
LW transient temporary
LB temporary
LR temporary
RR permanent temporary
ILC born on area permanent transient
LG born adjacent* permanent permanent
RBSH transient permanent
LPSS born on area temporary
STS born on area temporary
6404 temporary
72 born off area* transient temporaryt
6418 transient temporary
6477 temporary
6462 born adjacent # permanentt
6487 permanent
Total adults
Permanent 3 3 2 2
Temporary 2 3 4 3
*Immigrated on to study area as a juvenile.
t Alive in 1979.
#Immigrated on to study area as an adult in 1978.
IOVS
All losses of permanently resident males during the
summer were attributed to entry into hibernation
either because the males were recovered the following
year, or because they became progressively fatter and
less active, confining their movements to smaller areas
in the days immediately preceding their disappear-
ance.
After their emergence from hibernation, there were
always more resident females than males (Tables | and
2). Females did not exhibit a pattern of spring
movement and temporary residence, and all adult
females were resident on or near the study area, either
as juveniles or adults, in the previous year (Table 2).
Of the 11 original adult females resident in 1975, 10
remained for the summer, hibernated onthe area, and
reappeared in 1976. On 3 June a Badger (Taxidea
taxus) invaded the main burrow system of the 11th
female and she was not subsequently seen. A
neighboring, non-lactating female whose range was
adjacent to the study area extended her area to include
part of that formerly used by the missing female. This
female was not recovered in 1976.
Fourteen of the I5 females resident in 1976
remained through April and May and produced
litters. The missing female, when last captured on 8
April, had a scarred face, weighed 210 g compared
with an average of 276 + 29 g for eight other females
caught that day, and probably died in mid-April. Of
the remaining 14 residents, six were recovered in 1977
and must have hibernated on the area. Four other
females disappeared at a similar time to those that
were recovered and apparently hibernated, but did
not survive over winter. Four females disappeared in
June, too early to have gone into hibernation on the
area.
Of the 12 females resident in 1977, 11 remained on
TABLE 2—Composition of the population of adult females
resident on the study area each year
Number resident
Origin 1975 1976 1977 1978
Resident on area 1975 11 10 3 0
Born on area 1975 5 Ds |
Born on area 1976 6 3
Born on area 1977 0
Born adjacent 1975+ |
Resident adjacent 1976+ I l
Resident adjacent 1977+ I
Total permanent residents 11 IS) 12 i
*A third 2-yr-old was recovered but she established residency
adjacent to the study area.
+Females in these categories were captured at least once
within 40 m of the study area prior to the year they became
resident on the study area.
MICHENER: GROUND SQUIRREL POPULATION DYNAMICS
365
the area until the time of entry into hibernation, and
five of these were recovered in 1978. The 12th animal,
born on the area in 1975, disappeared in late May
when her pups were about 3 wk old. Death rather than
dispersal is the most likely explanation for her
disappearance during the lactation period. All seven
females resident in 1978 were still present when this
study ended on 24 April.
The number of adults hibernating on the area was
similar in each year: 3 males and 11 females in 1975, 3
males and 10 females in 1976, and 2 males and 11
females in 1977. Recovery rates in the following spring
varied: in 1976, 0 males and 10 females were recovered,
in 1977, 3 males and 6 females, and in 1978, 0 males and
5 females.
Juveniles
The numbers of adult female squirrels that re-
mained on the area throughout the breeding, gesta-
tion, and lactation periods were 10, 14, and 11 for
1975, 1976, and 1977, respectively. Of these 9, 14, and
8 respectively gave birth, and 6, 12, and 7 reared at
least one pup to weaning age. I attributed the
complete and partial losses of four litters in 1975 to
Long-tailed Weasel (Mustela frenata) predation be-
cause I saw a weasel hunting in the natal burrow
systems of the mothers of these litters. Two of the
mothers subsequently extended their ranges and spent
less time in the area of the natal burrow, behavior
patterns normally seen after the young are weaned
(Michener 1979a), and they reared no young. The
other two weaned only one young each. Such losses
contributed to the smaller size of emerged litters in
NOS:
Average litter size + SD (range; N)
OWS) 2s) ae\ 5) Cl=s2 G)
1976 4.4 + 1.5 (1-6; 11)
1977 4.0 + 0.6 (3-5; 7)
The average litter size at birth for females captured
when pregnant from the surrounding grassland in
1976 was 4.9 + 1.0 (3-6; 15) (Michener 1977),
suggesting a loss of 0.5 young per litter from field
litters in 1976 during the 30d between birth and
weaning. No attempt was made to estimate in-burrow
losses of the 1977 litters.
Because 1976 had both the largest number of
females rearing litters and the largest emerged litter
size, the number of juveniles reared on the area that
year was greater than in 1975 and 1977 (Figure !).
There were 48 juveniles (28 males and 20 females) in
the 11 of the 12 litters for which adequate data were
available, compared with 15(7 males and 8 females) in
1975 and 28 (13 males, 13 females, 2 unknown) in
1977.
In 1975 only one of the 15 juveniles born on the area
failed to remain resident until fall; on 14 July I killed a
366 THE CANADIAN FIELD-NATURALIST Vol. 93
11 1:3 1:3 16° 1S we2i 5.5 0) OH) 2-0 1977
i) 6 litters
rae es Chemie ats
10 E |
(0) LJ | ao BG T | i —
14 1:3 1-0 0-8 0-8 1 7-0 3 1976
30 11 litters
wm
Sie2d ine He
> born on
—)
=> 20 B area
Ba Mis a
7a
ws :]
S a
= 10
a fo H ae
= e
: |
a
foo) 0 Il [] [sat
0-9 1-3 4 14 1-4 14 16 9:0 1975
coc
= 6 litters
co 10
: age ieoml ad Eels jae
=m = ]
0 4 R= =
15 31 15 30 15 31 15 31 15 30 15
MAY JUNE SURIAY, AUGUST SEY OCT
FIGURE |. Numbers of juvenile males (left bar) and juvenile females (right bar) resident on the area in 1975, 1976, and 1977.
The sex ratio (males per female) is given above each pair of bars. The arrows indicate the time in each year at which
juveniles first began leaving the population to hibernate.
female that had severe myiasis of the neck and
shoulder due to Sarcophaga. citellivora maggot
infestation. The remaining juveniles disappeared in
mid- and late September (Figure 1) but only five
females and two males were subsequently recovered as
yearlings in 1976.
Of the 28 male and 20 female juveniles born on the
area in 1976 none of the 19 males and 9 females that
disappeared before 28 August was recovered in 1977. I
assume that these losses were due to death or dispersal
and that the pre-hibernation population consisted of
those juveniles still resident on 28 August. There was
only one accountable loss: I killed a juvenile male with
Sarcophaga citellivora myiasis of the leg on 12
August. The initial sex ratio at litter emergence in
1976 was male-biased (Figure 1), but male loss was
greater than female loss, particularly during July and
early August when males were lost ata rate of 0.31-d |
compared with 0.13-d' for females. By mid-August
the sex ratio was female-biased and remained so until
juveniles began entering hibernation. Of the 9 males
and I]! females resident after 28 August two and six,
respectively, were recovered in 1977.
Losses from the 28 juveniles that emerged on the
area in 1977 commenced with the disappearance of
four young, comprising an entire litter, between their
emergence on 25 May and my next visit on 29 May.
Because the mother remained resident at the same
location, discounting the possibility of removal of her
litter to a new burrow, and because juveniles 30 to 33d
old are unlikely to disperse, these young probably had
died. A further three males and seven females
NOS
disappeared by the end of July. One of these males
when last captured on 12 July had a swollen tongue,
showed labored breathing, weighed 160 g compared
with 223 + 28 g for nine other juvenile males captured
that day, and presumably died. Because none of the
juveniles born in 1977 was recovered as a yearling in
1978, I could not attribute disappearance in late
summer to immergence as opposed to death or
dispersal for any individual. Assuming that im-
mergence did not commence before mid-August, at
most one female and seven males hibernated on the
area. Thus there was a total loss of at least 6 males, 12
females, and 2 newly emerged young of undetermined
SeX.
Each year juveniles appeared that had not been
born to resident females. If they used a burrow system
on the area for at least 2 wk they were classed as
resident immigrants (Figure 1). There were more
immigrant juveniles in 1975, the year with the lowest
squirrel density, than in 1976 or 1977. Of the five
males and one female classed as immigrants in 1975,
three of the males were known to have been born to
females that were resident immediately adjacent to the
area. Their appearance on the area may therefore
reflect selective use of the mother’s total range rather
than true dispersal away from the natal area.
Inadequate information is available on the other
immigrants to determine whether any had moved over
a large distance. Transient juveniles were also seen
each year; some were caught several times over the
summer and were probably nearby residents making
excursions.
Causes of Loss
Disappearance of a ground squirrel could be due to
three factors: death, dispersal, or entry into hiber-
nation. Usually cause of loss could be accurately given
only for those squirrels which reappeared the next
spring; they left the active population and hibernated.
In this study, of the 32 adult females, 8 adult males, 22
juvenile females, and 30 juvenile males resident
(including immigrants) at the time of entry into
hibernation, 21 (66%), 3 (38%), 12 (55%), and 6 (20%),
respectively, were recovered the following year,
indicating that greater male than female loss occurred
between the time the first squirrel in each cohort
disappeared underground and the time the last
squirrel in each cohort reappeared in spring. I believe
that the majority of the missing animals died after they
left the active population to hibernate. Because some
males were already active in 1976 and 1977 when the
area was first visited in the spring, more males may
have survived the hibernation period but moved off
the area. A suspected, though not verified, source of
mortality during the hibernation phase was predation
by terrestrial predators, particular Badgers, on torpid
MICHENER: GROUND SQUIRREL POPULATION DYNAMICS
367
squirrels over the several-month period between
immergence and freezing of the soil. Extensive Badger
digging on the study area occurred between 13
September and 28 October 1975, and between 28
August and 10 October 1976. Such predation could be
age or sex selective if there are any differences in the
quality or depth of hibernacula used by adults and
juveniles or by males and females.
A possible source of mortality of adult males during
early spring was intraspecific fighting. Once females
emerged from hibernation the frequency and extent of
wounds on the lower back, legs, and shoulders of
males increased; I do not know whether the damage
was inflicted by other males or by females. Such
wounding associated with the weight loss males
normally undergo during the breeding season could
result in death, directly or through susceptibility to
disease and predation.
Of the 7 adult females, 22 juvenile females, 25
juvenile males, and 2 juveniles of unknown sex lost
during the active season, two were known deaths
(killed because of Sarcophaga citellivora infestation)
and three were suspected deaths due to Badger
predation or poor health. None of the remaining 5
adults and 48 juveniles that disappeared during the
summer was subsequently recovered and cause of
disappearance was not known for any of them.
Some losses to predators undoubtedly occurred. I
saw Long-tailed Weasels, Badgers, and Coyotes
(Canis latrans) on the study area on 19%, 11%, and
15% of the days I spent there and I found 0, 13, and 17
scats in 1975, 1976, and 1977 respectively. Twelve of
the 13 scats found in 1976 and 13 of the 17 found in
1977 contained the remains of Richardson’s Ground
Squirrels. Although these remains may not have come
from squirrels on the study area, they do indicate that
Richardson’s Ground Squirrels are a usual source of
prey for these predators. Red-tailed Hawks (Buteo
Jamaicensis), Swainson’s Hawks (B. swainsoni), and
Golden Eagles (Aquila chrysaetos) were often sighted
and Bald Eagles ( Haliaeetus leucocephalus) were seen
every spring during their northward migration. One
hawk pellet found on the area in 1976 contained the
remains of an adult squirrel. In 1976 one carcass and
in 1977 two carcasses were found on the area; there
were insufficient remains to verify the cause of death
or whether these were resident squirrels.
I assume that some of the losses over the active
season were due to dispersal but, with the exception of
four adult males that were not summer residents but
were captured on the area in two consecutive springs,
there were no verified cases of dispersal from the
area. Dispersal onto the area occurred but often
involved only a small-scale movement of a nearby
resident. Of 4 adult females, 3 juvenile females, and 10
juvenile males that moved onto the area 4, 0, and 3
368
respectively were known to have been previously
resident within 50 m of the area. Four of the seven
males classed as permanent residents in 1976, 1977,
and 1978 had not been resident the previous year and
thus had immigrated in spring.
Discussion
The adult sex ratio among Richardson’s Ground
Squirrels is biased in favor of females (Nellis 1969;
Sheppard 1972; Michener and Michener 1977) but
virtually all females are bred (Michener 1974),
indicating that males range over an area sufficient to
encounter three or four females during the breeding
season. Although Yeaton (1972) stated that male
Richardson’s Ground Squirrels established territories
in spring that encompass the burrows of three to five
females, implying exclusive access to those females, he
neither defined his use of the term territory nor his
method of determining territorial size and bound-
aries. From my observations of squirrels in early
spring of 1976 I found that the ranges (see Michener
1979a for details of calculating range size) of each of
the six males classed as temporary or permanent
residents overlapped not only with the ranges of at
least five females but also with the ranges of at least two
males, while the ranges of the females overlapped the
ranges of between two and five males. Apparently no
male had exclusive access to any female. Because
females are bred soon after emergence in spring,
exclusive access may occur if males make day-to-day
variations in their ranges relative to the location of
receptive females.
This study indicated that in spring males remained
on the study area for periods ranging froma few hours
to several days to the entire season. Because of the
presence of transients and temporarily resident males,
more males were trapped in early spring than
subsequently established residency within the area. If
males had not been classed according to their status
on the area, trapping data alone would have revealed
male loss in spring suggesting differential mortality
favoring females, as reported by Dorrance (1974) and
Schmutz et al. (1979). The size of the ranges occupied
by males declined after the breeding season (Michener
1979a) because of a decrease in the length and
frequency of movements made by males. Although
some loss of males in spring was probably due to
mortality, the ‘loss’ of at least seven males that were
transients or temporary residents was related to this
decrease in movements resulting in a smaller range
that no longer overlapped the study area. By the time
females began giving birth no further changes in range
usage by males occurred and all males still present
remained as summer residents. Adult males that had
been born on the area also changed their residency
status with respect to the study area within any one
THE CANADIAN FIELD-NATURALIST
Vol. 93
breeding season and between consecutive breeding
seasons. Because female Richardson’s Ground Squir-
rels remain in the same area over several years one
effect of male movement in spring is to reduce the
amount of inbreeding.
This study confirmed previous observations (Yea-
ton 1972; Michener 1979a) that adult females, .
including yearlings, make only small-scale changes in
the location of their ranges between years and within
years. Female losses from the area during the active
season were low in 1975 and 1977, and were attributed
to mortality. In 1976, however, four adult females
disappeared in June. The period immediately after
lactation would be most suitable for dispersal of adult
females; dispersal during the breeding, gestation, or
lactation periods would be disadvantageous to suc-
cessful rearing of a litter, and dispersal later in the -
summer would interfere with the weight gain that
follows weaning of the litter and might not allow
sufficient time to locate a suitable hibernation
burrow. This study and those of Dorrance (1974) and
Michener and Michener (1977), however, have found
no evidence for dispersal by adult females. The higher
recovery rate of females hibernating over 1975-1976
(91%) compared with 1976-1977 (60%) and 1977-
1978 (45%) may be related to the higher proportion of
females raising no young or only one young in 1975.
Weather conditions unsuitable for feeding in the late
summer of 1976 and the more severe winter over
1977-1978 (Michener 1979b), combined with the
demands of rearing larger litters in 1976 and 1977,
probably contributed to the reduced survival in the
1976-1977 and 1977-1978 winters. Whatever the
causes, mortality during the overwinter period does
appear to be the main factor affecting the size of the
population of adult female Richardson’s Ground
Squirrels.
Each year fewer juveniles emerged than expected on
the basis of the reproductive potential of the popula-
tion, and there was greater between-year variation in
the size of the juvenile population at emergence than
expected from the differences in the number of adult
females resident each spring. Normally 100% of
yearlings and older females are successfully bred, and
between 90% and 100% lactate (Nellis 1969; Sheppard
1972; Michener 1974; this study in 1975, 1976, and
1978): the unexplained failure of three yearlings to
produce litters in 1977 reduced the potential juvenile
population by about 12. Comparison of litter size at
birth (Michener 1977) or in utero (Schmutz 1977) with
size of field litters at emergence indicates that there is
usually a small loss from most litters. Predation on
litters by weasels, however, caused major losses and
largely accounted for the appearance of only 15
juveniles in 1975 when about 36 were expected from
the number of resident lactating females.
1979
There was a net incease of juveniles between
emergence from the natal burrow and the time of
entry into hibernation in 1975 but net loss in the other
two years. In 1975 no juveniles left the area, suggesting
that there are densities below which dispersal does not
occur, but the immigration rate was highest, indicat-
ing that dispersal can affect the size of the juvenile
population. Although indirect evidence suggests that
terrestrial predators took a larger number of squirrels
in the years with the highest squirrel densities, | could
not estimate the magnitude of losses or determine
whether such losses were sex-related. Luttich et al.
(1970) reported that the sex ratio of juvenile Richard-
son’s Ground Squirrels killed by Red-tailed Hawks
approached equality, whereas Schmutz et al. (1979)
found that Swainson’s Hawks and Ferruginous
Hawks (Buteo regalis) captured disproportionately
more juvenile males. I believe that a combination of
dispersal and predation accounted for the majority of
losses of juvenile males in 1976 and 1977. Male loss
exceeding female loss, as in 1976, has previously been
reported for juvenile Richardson’s Ground Squirrels
(Dorrance 1974; Schmutz et al. 1979) and other
hibernating ground squirrels (Rongstad 1965;
McCarley 1966: Slade and Balph 1974: Morton et al.
1974: Dunford 1977). Greater loss of females is
unusual and I do not know why 90% of juvenile
females disappeared in the summer of 1977. With the
exception of females in 1977 the major loss of
juveniles from the area and the major immigration of
juveniles onto the area occurred during July, as in the
studies by Dorrance (1974) and Schmutz (1977),
suggesting that there is a minimum age juveniles attain
before dispersing and that dispersal is completed at
least a month before hibernation commences.
Although the sex ratio (males per female) among
juveniles in the pre-hibernation population varied
from 1.4 in 1975, to 0.8 in 1976, to 9.0 in 1977, the sex
ratio among yearlings recovered was female-biased in
1976 (0.6) and 1977 (0.4); with no recoveries in 1978,
suggesting greater loss of males than females over-
winter. Mortality occurring at the end of the active
season, however, when juvenile males were predomin-
ant in the active population, or at the beginning of the
next active season, when adult males predominantly
constituted the active population, would also affect
the sex ratio. Schmutz et al. (1979) considered that
mortality, mainly due to predation by migrating avian
predators, at these times was significant in readjusting
the sex ratio, whereas Michener and Michener (1977)
believed overwinter mortality to be more important in
the readjustment. Laboratory studies of hibernating
sciurids show that there are basic differences between
males and females, with males having lower survival
rates, longer bouts of continuous torpor but a shorter
hibernation season, and a greater likelihood of
MICHENER: GROUND SQUIRREL POPULATION DYNAMICS
369
permanently arousing when stressed or disturbed
during hibernation (Twente and Twente 1967; Morri-
son and Galster 1975; Pengelley et al. 1978), so higher
mortality of males during the hibernation phase is a
possibility.
As with previously reported studies (Dorrance
1974: Michener and Michener 1977; Schmutz et al.
1979) of Richardson’s Ground Squirrels I found that
there was high inter-year loss of squirrels, with male
losses exceeding female losses and juvenile losses
exceeding adult losses, and that adult female squirrels
rarely dispersed, with the major loss from this cohort
occurring over winter. Here I have indicated that
observation or trapping at frequent intervals during
early spring is necessary to reveal the residency status
of adult males. The contraction in size of range used
by a male once all females are bred contributes to the
apparent loss of adult males from a local area. Some
losses result, not from dispersal or predation, but
from the male confining his activity to a smaller range
outside the study area and thus being overlooked. |
found, as did Dorrance (1974), that variations in the
overwinter survival of females and in the extent of loss
of pups prior to weaning determined the density of the
population in mid-summer. Michener and Michener
(1977) predicted that when the adult female popula-
tion is small or the reproductive success of females is
low, there will be no juvenile dispersal. In this study
weasel predation, reducing juvenile addition to the
population, resulted in the lack of dispersal of
juveniles from the area in 1975. The size of the
population of adults in spring and of juveniles in
summer varied each year as a result of these interac-
tions between overwinter survival, reproductive suc-
cess, predation, and dispersal.
Discrepancies among Dorrance (1974), Michener
and Michener (1977), Schmutz et al. (1979), and this
study in accounting for the greater losses of both adult
and juvenile males compared with females can, in
part, be attributed to variations in the populations
under investigation. Populations located in aspen
parkland (Dorrance 1974) and on short-grass prairie
in Saskatchewan (Michener and Michener 1977), in
Alberta (Schmutz et al. 1979), and at the western limit
of the species’ range (this study) have been studied for
2,3, or 4 continuous years between 1969 and 1978, and
the densities of adults have ranged from about | per ha
(Michener and Michener 1977) to 10 or more per ha
(Dorrance 1974). If, as seems likely, a combination of
dispersal, predation, and overwinter mortality poten-
tially contribute to total losses and to differential
losses among sexes, then in any given location in any
year one factor may predominate without eliminating
the others as important factors under other condi-
tions. Disputes between the relative importance of
predation and overwinter mortality currently revolve
370
around persuasive arguments rather than empirical
evidence, and will only be resolved by studies in which
every individual is traced, by telemetry, to distinguish
entry underground from late-season mortality and to
distinguish death underground from early emergence
in spring followed by death or dispersal.
Acknowledgments
The first two years of this study were conducted
while I held an Isaac Walton Killam Memorial Post-
doctoral Fellowship at the University of Alberta.
Support and equipment were provided by National
Research Council of Canada grants to A. L. Steiner
and J. O. Murie of the University of Alberta. J. O.
Murie and D. R. Michener reviewed the manuscript. I
thank L.S. Davis for providing information on
survival of males in 1979, and Mr. and Mrs. J. Bews of
the Y-Cross Ranch Ltd. for permitting access to their
land.
Literature Cited
Dorrance, M. J. 1974. The annual cycle and population
dynamics of Richardson’s Ground Squirrels. Ph.D. thesis,
University of Wisconsin, Madison. 150 pp.
Dunford, C. 1977. Behavioral limitation of Round-tailed
Ground Squirrel density. Ecology 58: 1254-1268.
Luttich, S., D. H. Rusch, E. C. Meslow, and L. B. Keith.
1970. Ecology of Red-tailed Hawk predation in southern
Alberta. Ecology 51: 190-203.
McCarley, H. 1966. Annual cycle, population dynamics
and adaptive behavior of Citellus tridecemlineatus. Jour-
nal of Mammalogy 47: 294-316.
Michener, D. R. 1974. Annual cycle of activity and weight
changes in Richardson’s Ground Squirrel, Spermophilus
richardsonii. Canadian Field-Naturalist 88: 409-413.
Michener, G. R. 1977. Effect of climatic conditions on the
annual activity and hibernation cycle of Richardson’s
Ground Squirrels and Columbian Ground Squirrels. Can-
adian Journal of Zoology 55: 693-703.
Michener, G. R. 1979a. Spatial relationships and social
organization of adult Richardson’s Ground Squirrels.
Canadian Journal of Zoology 57: 125-139.
Michener, G. R. 1979b. The circannual cycle of Richard-
son’s Ground Squirrels in southern Alberta. Journal of
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Mammalogy 60(4). /n press.
Michener, G.R. and D.R. Michener. 1977. Population
structure and dispersal in Richardson’s Ground Squirrels.
Ecology 58: 359-368.
Morrison, P. and W. Galster. 1975. Patterns of hibernation
in the Arctic Ground Squirrel. Canadian Journal of
Zoology 53: 1345-1355.
Morton, M.L., C.S. Maxwell, and C.E. Wade. 1974.
Body size, body composition, and behavior of juvenile
Belding Ground Squirrels. Great Basin Naturalist 34:
121-134.
Nellis, C. H. 1969. Productivity of Richardson’s Ground
Squirrels near Rochester, Alberta. Canadian Field-
Naturalist 83: 246-250.
Pengelley, E. T., R. C. Aloia, B. Barnes, and D. Whitson.
1978. Comparative temporal aspects in hibernation be-
tween male and female Golden-mantled Ground Squirrels,
Citellus lateralis. Journal of Thermal Biology 3: 88.
Rongstad, O. J. 1965. A life history study of Thirteen-lined
Ground Squirrels in southern Wisconsin. Journal of
Mammalogy 46: 76-87.
Schmutz, S. M. 1977. Role of dispersal and mortality in the
differential survival of male and female Richardson’s
Ground Squirrels. M.Sc. thesis, University of Alberta,
Edmonton. 97 pp.
Schmutz, S.M., D. A. Boag, and J. K. Schmutz. 1979.
Causes of the unequal sex ratio in populations of adult
Richardson’s Ground Squirrels. Canadian Journal of
Zoology. /n press.
Sheppard, D.H. 1972. Reproduction of Richardson’s
Ground Squirrel (Spermophilus richardsonii) 1n southern
Saskatchewan. Canadian Journal of Zoology 50: 1577-
1581.
Slade, N. A.and D. F. Balph. 1974. Population ecology of
Uinta Ground Squirrels. Ecology 55: 989-1003.
Twente, J. W. and J. A. Twente. 1967. Seasonal variation
in the hibernating behavior of Citellus lateralis. In Mam-
malian hibernation III. Edited by K.C. Fisher, A. R.
Dawe, C. P. Lyman, E. Schonbaum, and F. E. South.
Elsevier, New York. pp. 47-83.
Yeaton, R.I. 1972. Social behavior and social organiza-
tion in Richardson’s Ground Squirrel (Spermophilus
richardsonii) in southern Saskatchewan. Journal of Mam-
malogy 53: 139-147.
Received 24 February 1979
Accepted 30 May 1979
Man?’s Influence on Potential Nesting Sites and Populations of
Swallows in Canada
A. J. ERSKINE
Canadian Wildlife Service, Sackville, New Brunswick. E0OA 3CO0O
Erskine, Anthony J. 1979. Man’s influence on potential nesting sites and populations of swallows in Canada. Canadian
Field-Naturalist 93(4): 371-377.
The nesting of the seven species of swallows (Hirundinidae) occurring in Canada was examined using the nest records
assembled in Canadian nest records schemes. All species have been influenced by the actions of modern man, and more have
been favourably affected through increased availability of nest sites than have declined, despite adverse competition from
exotics introduced by man. Changes in breeding range related to environmental changes caused by man have occurred and are
still occurring.
On a examiné la nidification de sept espéces d’hirondelles (Hirundinidae) au Canada, utilisant des fiches rassemblées dans les
fichiers de nidification des oiseaux canadiens. Les activités de homme depuis la colonisation européene ont influencé toutes
les sept espéces, mais la plupart des espéces ont profité a cause de la disponibilité augmenteée des sites de nidification. Les
alterations de l'environnement causée par "homme ont aussi influence la distribution de quelques espéces d’hirondelles, et
tout ¢a continue toujours.
Key Words: Hirundinidae, swallows, nesting sites, Canada, nest records, populations.
Seven species of swallows (Hirundinidae) are
widespread in Canada and the United States, includ-
ing two which also breed in Europe, Asia, and North
Africa, and another which breeds also in South
America. All are at least moderately gregarious, and
several nest colonially. Nesting is restricted more to
particular situations, sites, or substrates than to
special habitats. This paper explores some of the
effects of actions by man, particularly since white
settlement, on availability of nesting sites for North
American swallows, and speculates on their indirect
effects on numbers of those birds.
Materials and Methods
I examined all records through 1974 of swallow
nests in the five major Canadian nest records schemes
(NRS) (see Acknowledgments). The few records (less
than 20 per species) in the Newfoundland NRS were
omitted. Each nest was categorized as in a natural site
or one affected by man’s actions; some were not
certainly assignable to either category. Obvious
duplication within a year was eliminated, but not that
between years. Published summaries (e.g., Bent 1942:
Graber et al. 1972) and authoritative works on each
species (q.v.) were consulted. Those sources described
the entire spectrum of nest sites reported on the nest
record cards without suggesting that any important
types had been missed, and consequently no
exhaustive literature review was attempted.
The relative abundance of each species in different
parts of its range in Canada was inferred from results
of the co-operative Breeding Bird Survey (BBS)
(Erskine 1978, and unpublished data). Those data are
believed suitable for broad comparisons of relative
density within a species, but not for comparisons
between species.
Results
Data for each species are summarized below.
The species are arranged according to type and
construction of nests, rather than in taxonomic order.
The sequence followed starts with species that use
existing cavities, first those that were originally (as far
as we can infer) obligate tree-hole nesters, followed by
more tolerant species that accept cliff crevices and
bank burrows as well as tree holes. Last come species
that actively prepare their own nests by excavating
burrows or by building fully or partly enclosed
structures from clay pellets.
PURPLE MARTIN (Progne subis)
Our largest swallow is the northernmost represen-
tative of a South American group. Its history of
association with man extends back before white
settlement, when martins already nested in gourds
hung up for them by Amerindians (reviews by Allen
and Nice 1952: Jackson and Tate 1974). Its use of
natural nest sites is now seldom reported in the east:
presumably those were in tree cavities, with no more
than four or five nests in any tree, as was still the case
recently in the west (Richmond 1953; Finlay 1975).
Among roughly 3000 nests reported by Canadian nest
records schemes, only seven were stated to be in trees
or stubs, and five of those were from British
Columbia’s small coastal population, from which
only one nest box record was also available.
Now most martins use artificial nest boxes, includ-
ing multiple-unit “hotels” as well as single boxes.
Sil
S72
Many Canadian nest records reported only the
number of “hotels,” which may have up to 30 or more
compartments; in such cases the number of occupied
nests could only be guessed at. Mean colony size,
calculated from reported data, varied from five pairs
in Ontario to nine in New Brunswick, ranging from
isolated pairs up to clusters of hotels housing over 50
pairs. In Canada, martins are found mostly in towns
and cities, and 24 old records from Ontario referred to
nests within the walls or roofs of buildings.
TREE SWALLOW (Jridoprocne bicolor)
The arrival each year of this relatively boreal
swallow represents “spring” to many Canadians. It
nests singly in tree holes and nest boxes, often near
human habitations. Its natural sites are relatively
difficult to find and to inspect; only 14% of 4370
reported sites were in tree holes, largely of wood-
pecker origin (Table |). Natural sites made up an
appreciable fraction of the available records only in
British Columbia (Table 1), where several thesis
studies had concerned themselves with hole-nesting
birds (Erskine 1960; McLaren 1963: Kelleher 1963).
The vastly higher proportion of natural sites in British
Columbia may also reflect the relatively higher
density there of flickers (Co/aptes) (cf., Erskine 1978,
Tables 3-8), whose nest cavities are often used by Tree
Swallows.
There were only two records, both by experienced
observers, of Tree Swallows nesting in holes in earth
or rock cliffs; cavities in the walls of buildings were
seldom used by this species. Tree Swallows evidently
find cliff sites unacceptable, and should be viewed as
having been obligate tree-hole nesters originally,
unlike the following species.
Most nest sites classified as affected by man’s
actions were nest boxes, usually erected for bluebirds
(Sialia spp.), diving ducks (Bucephala spp.), or for
Tree Swallows specifically. Cavities excavated by
woodpeckers in poles or posts erected by man were
also listed as affected by man.
THE CANADIAN FIELD-NATURALIST
Vol. 93
VIOLET-GREEN SWALLOW
(Tachycineta thalassina)
A species confined to western North America, this
swallow either breeds as single pairs in tree cavities,
openings within the walls or roofs of buildings, or in
artificial nest boxes, or else in groups in rock crevices
(Edson 1943). All of the 550 Canadian nest records
were from British Columbia, and 42% were in natural
sites, chiefly as colonies in cliffs. Nest boxes and the
walls and eaves of buildings accounted for most of the
other 58%.
Cliff-nesting might be expected to have been
originally more prevalent in the sparsely vegetated
interior areas of British Columbia, with tree cavities
predominating in the coast forests. Unfortunately, I
did not segregate the different types of nest sites used
by geographic area within the province. We do not
know whether birds reared in tree holes will accept
holes in buildings (artificial cliffs) or only nest boxes
(artificial tree holes), and conversely whether birds
reared in cliff sites will or will not accept other sites.
ROUGH-WINGED SWALLOW
(Stelgidopteryx ruficollis)
This brown-backed swallow breeds from South
America to southern Canada, and most nest records
examined were from southern Ontario and British
Columbia. It nests in small groups and as isolated
pairs in various bank and cliff situations, using
burrows (some or all made by other species) (Lunk
1962) in earth banks, or existing openings in rock
walls. Excluding unassignable records, 68% of the 725
nests reported were in natural sites and only 32% in
burrows in gravel pits or road cuttings, or (especially
in Ontario) holes in cement, brick, or stone walls (cf..,
Lewis 1944).
Whereas burrows by rivers predominated in British
Columbia, man-made sites were most commonly used
in Ontario, the most settled (by humans) and altered
part of Canada (Table 2). Colony size was higher in
British Columbia than elsewhere (Table 2), and the
TABLE 1—Regional variations in proportions of natural and man-made sites used by Tree Swallows in Canada
% of recorded nests in
Natural sites
Man-affected sites
Woodpecker holes Woodpecker holes Nest Other
Region in trees and stubs in poles and posts boxes sites Total nests
Maritime provinces 4 5) 88 2 466
Quebec and Ontario 9 4 86 1 2113
Prairie provinces 4 | 93 2 1059
British Columbia 46 4 44 6 752
All regions 14 3 81 2 4370
1979
ERSKINE: CHANGES IN SWALLOW NESTING SITES
Sif)
TABLE 2—Regional variations in proportions of natural and man-made sites used by Rough-winged Swallows in Canada
% of assignable nests
Uncategorized
Natural Man-made Total number nests
Region sites sites (mean number/ colony) (% of gross)
Maritime provinces 100 0 I (1) 0
Quebec and Ontario 50 50 213 (2) 19
Prairie provinces 78 22 18 (2) 0)
British Columbia US DS 493 (15) 12
All regions 68 32 125 (4) 14
species is also relatively more common in that
province, judged by BBS data.
BANK SWALLOW ( Riparia riparia)
In Eurasia as well as North America, this small
swallow nests colonially in burrows which it excavates
in near-vertical banks of earth, clay, and fine gravel
(Petersen 1955). Nearly one-third of the nest records
could not be categorized as natural or affected by
man. Of those assigned (27718 in all) about 40% were
in natural sites, mainly along sea coasts, rivers, and
lake shores where erosion by water or wind led to
exposure of earth slopes. The remaining 60% were in
cut banks created by man — sand and gravel pits, road
and railway cuttings, and various excavations, except
for about 1% in piles of gravel stored for road repairs
(cf., Nero 1968), and a few in old sawdust heaps (cf.,
Greenlaw 1972).
The data for various regions of Canada are
summarized in Table 3. Natural sites were more often
reported where human densities are relatively lower,
and particularly in the Maritimes where coastal cliffs
are also frequent. Man-made sites made up a larger
proportion of the records in Quebec and Ontario,
where human population, and presumably also
habitat disturbance, is greatest. Man-made situations
are predominant in British Columbia, where most
records reported nests in roadside cut banks. Munro’s
monograph (1945) on birds of the Cariboo Parklands,
then an undeveloped area with rather primitive roads,
omitted Bank Swallows, which now are regular there
(Erskine and Stein 1964). Road-building may thus
have permitted more general distribution of a
formerly localized species.
Mean colony size was much lower in the prairies
than elsewhere. The relative paucity of nest records as
well as the relative density data from BBS suggest that
Bank Swallows are scarcer there too.
CLIFF SWALLOw ( Petrochelidon pyrrhonota)
Cliff Swallows build up their nests from clay pellets
plastered against vertical or overhanging walls of cliffs
or structures. The nests are fully enclosed, gourd-
shaped structures, often assembled in large colonies
(Mayhew 1958). Such nests depend on virtually
complete overhead protection from rain and runoff
water, whether on cliffs or buildings. Of 21096
Canadian nest records, only 23% were in natural sites
on cliffs, the rest being on buildings, bridges, culverts,
and dams.
Cliffs are presumed to be more prevalent in the
western mountains than in most of the flatter regions
farther east, and records of natural cliff nest sites were
almost all from the west (Table 4), with colonies of
1000 or more nests along the Bow River near Calgary,
Alberta. The latter biased the regional comparisons
(Table 4) of natural vs. man-affected sites and of mean
colony size, as no comparably large colonies were:
reported in British Columbia. The BBS density
indices, however, were as high for those parts of
British Columbia where the species was recorded as
for Alberta.
TABLE 3—Regional variations in proportions of natural and man-made sites used by Bank Swallows in Canada
% of assignable nests
Uncategorized
Natural ~ Man-made Total number nests
Region sites sites (mean number/colony) (% of gross)
Maritime provinces iS 25 8207 (56) l
Quebec and Ontario 35 65 9934 (38) 39
Prairie provinces 57/ 43 509 (5) 0
British Columbia 13 87 8568 (59) 35
All regions 40 60 27218 (42) 29
374
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 4—Regional variations in proportions of natural and man-made sites used by Cliff Swallows in Canada
% of recorded nests in
Total number
Natural Man-made of nests
Region sites sites (mean number/ colony)
Maritime provinces trace* 100- 1305 (19)
Quebec and Ontario trace* 100- 1702 (8)
Prairie provinces 57 43 6735 (102)
British Columbiat 9 91 11354 (31)
All regions 23 V7) 21096 (30)
*Trace = much less than 0.5%.
tIncluding a few from Yukon Territory and Northwest Territories.
BARN SWALLOW (Hirundo rustica)
This composition species is “the swallow” of
English proverbs, as familiar in Eurasia as in North
America. Barn Swallows build their bracket-shaped
mud-pellet nests solitarily or in loose colonies inside
barns and other open buildings and under the porches
and eaves of occupied homes (Samuel 1971), although
the original sites were presumably all on overhanging
cliffs or in caves. Of the eight locations in Canada
where natural sites were reported in use, all in the
eastern provinces, four were in national parks and two
in a provincial park. Possibly such parks are among
the few areas frequented regularly by man where
buildings are scarce or lacking. Among nearly 5000
nests recorded in Canada, only 48 (1%) were on cliffs
or in caves, the rest being in (54%) or on the outside of
buildings (33%), or on other structures (12%).
Perhaps more than any of our other swallows, its
range may be still changing rapidly in response to
man’s activities, as is discussed later.
Discussion
(1) Biases. The nest records reflect the sites that
observers were able to find and inspect. Those are
certainly biased towards man-altered habitats and
man-associated nest sites (cf., Erskine 1971), so that
nests located by roadsides, around houses and
settlements, and in agricultural areas are certainly
reported in larger proportions than the natural sites in
undeveloped, and unvisited, areas. When the vast
majority (over 99%) of sites reported are man-
influenced, however, as for Purple Martins and Barn
Swallows, this is likely to reflect the real situation.
(ii) Effects of man on natural nest sites of swallows.
Man’s impact on natural nest sites probably affected
tree-hole nesters most. At the time of European
settlement, eastern Canada south of the tundra was
almost entirely forested. Subsequently, southern
Ontario and the St. Lawrence lowlands of Quebec, as
well as parts of the Maritime Provinces, were largely
cleared for settlement and agriculture (cf., Edwards
1969). Similar clearing of formerly forested land took
place all along the northern edge of the prairies, and in
the lower Fraser Valley of British Columbia. Every-
where, the large trees were the first to go. More
recently, dead trees and snags are being eliminated, on
the grounds of unsightliness in settled areas, and as
foci of infection by forest insects or as fire hazards in
cutover and regenerating forest lands. Thus, the large
and dead trees in which woodpeckers most often make
cavities — usable by swallows — have been reduced
much more than the present extent of forest lands
might suggest. Man has probably had little overall
influence on availability of other types of natural nest
sites used by swallows.
(iii) Trends in use of new nest sites resulting from
man’s activities. Despite man’s effect in reducing
availability of natural sites for some swallows, man-
made situations undoubtedly have provided many
additional nesting opportunities. Purple Martins are
perhaps the most “domestic” of swallows now, as
most of the sites they use are deliberately provided for
them by man, in close proximity to human dwellings.
This association began before white settlement, and
adaptation to artificial sites may have been gradual.
Although martins often assemble in far larger colonies
in man-made boxes than can have existed in natural
tree sites, the size of colonies still averages only 5.8
nests (calculated from data in Jackson and Tate 1974).
Tree Swallows were originally no less obligate as
tree-hole nesters than martins, but their dependence
on man-made sites is much less. Possibly their
association with man is of more recent origin, since
their more northern distribution would have brought
them in contact, before white settlement, mainly with
Amerindians of nomadic habits, whereas the early
“adoption” of martins was by sedentary agricultural
tribes (Allen and Nice 1952).
The importance of nest boxes to Tree Swallows can
easily be overestimated. I suggest, largely on the basis
of “bluebird trail” reports, that nest boxes systemati-
1979
cally erected for bluebirds and other birds may
accommodate perhaps 10000 pairs of Tree Swallows
annually across Canada, and backyard nest boxes
may add another 10000 pairs, say 20000 pairs in all.
This looks impressive, but I have also estimated, from
density data given by Graber and Graber (1963) and
Stewart and Kantrud (1972) and the BBS density
indices (Erskine 1978, Tables 3-8), that the total
Canadian population of Tree Swallows may be
roughly one million pairs. Neither of these estimates is
precise, but they should serve to provide the necessary
perspective. The increase effected by use of nest boxes
is apparently in the order of 2%, which is almost
certainly insufficient to counterbalance the losses of
natural tree nest sites throughout the settled parts of
Canada in the last 200 yr.
The apparent ability of Violet-green Swallows to
use both tree and cliff sites deserves study. That
species is tolerant of a wide spectrum of nesting
cavities, which presumably has eased its acceptance of
man-made sites. Its domesticity in using backyard
nest boxes and holes in the eaves of houses is in
marked contrast to its nesting in remote cliffs. Rough-
winged Swallows, which also use crevices in rock
cliffs, are possibly the least domestic of all Canadian
swallows: they have made no real adaptation in using
existing cavities in man-made as well as natural banks
and cliffs. It seems likely that man has not had much
influence on nesting by those two species.
Bank Swallows excavate their burrows in man-
made banks exactly as they do in natural situations.
The much greater availability of cut banks and
excavations, even in flat areas, undoubtedly allows
more general distribution of that species than in
former times.
Nesting sites for Cliff Swallows have become far
more generally available as a result of man’s actions.
That species is much more widespread and abundant in
the west than ineastern North America (Erskine 1978,
Tables 3-8), as was also the case in the early 1800s
(Bent 1942). In contrast, Barn Swallows have an eastern
predominance today (Erskine 1978), but natural sites
cannot originally have been numerous /in eastern
North America, as is also suggested by the present
scarcity of the species in areas without human
settlement. The readiness of Barn Swallows to build
nests on newly-constructed buildings suggests that
they may have colonized permanent dwellings as soon
as these were built by European settlers in America.
Barn Swallows have recently spread southwards in
the southeastern United States, as shown by
comparison of recent BBS data with the range given in
the AOU Checklist (1957) only 15 yr earlier (D.
Bystrak, United States Fish and Wildlife Service,
unpublished data; Reid 1975). Prior to 1964, the Barn
ERSKINE: CHANGES IN SWALLOW NESTING SITES
3/5
Swallow also had not been recorded in northern
Alberta, an area with no cliffs and few settlements
(Erskine 1968), but with opening of that area for
agriculture and oil prospecting it is now seen
regularly. Samuel (1971) thought there was little
competition between Barn and Cliff Swallows in West
Virginia, although they frequently occurred together.
(iv) Effects on swallows of competition by introduced
exotics. The introduction to America by European
settlers of the House Sparrow (Passer domesticus)
and Starling (Sturnus vulgaris) has had adverse effects
on most native hole-nesters with which the newcomers
compete for nest sites. In the southern states, where
Purple Martins are relatively more common and
Starlings and sparrows less common than farther
north, Jackson and Tate (1974) considered such
competition a minor influence on martin numbers.
But farther north, near the limits of their range, some
martin colonies have been abandoned as a result of
competition with sparrows and Starlings when no
efforts were made by man to protect them (e.g.,
Brinkman, in Graber et al. 1972). In the west (Oregon
and Alberta), Purple Martins did not accept nest
boxes until after the introduced competitors had
arrived. House Sparrows were extremely local on the
west coast until 1900-1910 (Robbins 1973), and
Starlings did not breed in British Columbia until
about 1948 (Myres 1958).
One or other of the Tree or Violet-green Swallow
was the most common bird species in three northern
frontier towns in which both House Sparrows and
Starlings were present (Erskine 1977, Table 12), but
census data show that in southern Canadian cities House
Sparrows and Starlings are much more common than
any swallow species (Speirs et al. 1970; Weber 1972).
Detailed studies of competition are lacking, but
competition with the introduced pests has presumably
influenced both Violet-green and Tree Swallows,
especially in urban areas.
Bent (1942) quoted a few records of House
Sparrows and Starlings using Bank Swallow burrows,
and this is evidently rare in Canada too. Competition
from House Sparrows adversely affects Cliff Swal-
lows nesting on farm buildings (Samuel 1969), but
probably has less influence on those nesting in other
situations. It seems likely that the rapid initial spread
in range and increase in numbers of Cliff Swallows in
the east, following their acceptance of buildings as
nesting situations in the early 1800s (Bent 1942), was
reversed less than a century later by the influence of
House Sparrows. Barn Swallows are very seldom
bothered by House Sparrows, which prefer a fully
enclosed nest site.
(v) Effects on swallows of possible changes in insect
populations. In the Maritime Provinces and probably
376
also in eastern Ontario, I have the impression that
Tree Swallows have become markedly scarcer over
the past 25 yr, and especially between 1950 and 1960,
both absolutely and relative to Bank and Barn
Swallows (cf., my comments in Boyer 1966). The low
numbers I found in the Maritimes in 1960 could have
reflected losses in 1958 and 1959, but that leaves
unexplained their failure to recover subsequently, and
their similar scarcity in eastern Ontario after | moved
there in 1968. Competition with House Sparrows and
Starlings is unlikely to be implicated, as both exotics
were well established in those areas before 1950. I
believe the decrease more likely to reflect a reduction
in flying (and biting) insects in and around urban
areas as a result of the widespread use of DDT and
other insecticides for forest spraying and mosquito
control. Such insects were still conspicuously present
in the frontier towns (with abundant swallows) that I
censused in 1970-1975, although I have no quant-
itative data on insects. Increased use of cars, with
resultant increase in air pollution, is also well
correlated in time and space with the apparent recent
decline of Tree Swallows in urban areas.
The continued relative success of Purple Martins in
urban areas may reflect their tendency to forage
higher above the ground than other swallows. Local
insecticide spraying and air pollution in cities and
towns may have less effect in reducing the high-flying
insects on which martins feed.
(vi) Conclusions. Historical records generally pro-
vided little information other than presence inan area
as opposed to absence from it, which is of use in
assessing changes in distribution, but little help in
detecting trends in numbers. The changes in numbers
which I am suggesting in Table 5 are based on my
experience in using nest records and in interpreting
THE CANADIAN FIELD-NATURALIST
Vol. 93
population data, and on my personal impressions
from 25 yr of field work across Canada. My
suggestions are certainly speculative, and some people
will question my summation of the various influences
acting on swallows.
The order in which the species were discussed
reflected their success in coping with man’s effects on
their environment as well as the types of nests they
used. The obligate tree-hole nesters, discussed first,
depended on trees large enough to accommodate
cavities. Those birds seem to have come off least well,
losing potential nest sites with the cutting of trees for
settlement and agriculture as well as for timber, and
suffering from competition by introduced hole-
nesting species. The more tolerant cavity-nesting
swallows have been able to cope better with man-
caused changes, but have neither decreased nor
increased. Finally, the species that actively construct
their own nests have been able to take advantage of
either man’s penchant for excavation or his buildings,
and those species all have prospered. None of our
swallows, however, can be considered extreme ex-
amples of response to man’s actions; none has been
reduced to threatened status or extirpated, either
through direct persecution or damage to its environ-
ment; and none has burgeoned to the point of
becoming a pest. Few other families of passerine birds
have become as closely associated with man without
achieving either domestic or pest status.
I believe it desirable to show that not all species of
birds are necessarily affected adversely by changes
brought on by man. Even highly migratory and
esthetically pleasing species, including several of the
swallows, have become almost totally dependent on
man for nest sites. We should be aware of the extent of
such dependence when we make environmental
decisions.
TABLE 5—Man’s effects on swallow numbers in Canada, judged from nest-site selection and availability. Key to impacts:
— major decrease; — moderate decrease; 0 little or no change; + moderate increase; ++ major increase
Impact of man’s actions on swallows
Availability
of natural
Species sites
Purple Martin =
Tree Swallow —
Violet-green Swallow 0; =]
Rough-winged Swallow (0)
Bank Swallow (0)
Cliff Swallow 0)
Barn Swallow oO
Suggested
changes from
original
Status
Availability
of new sites
Competitors
introduced*
Qe | 2 ©
+++
+44
t++oo) |
*House Sparrow, Starling.
+Cliffs and trees listed separately, also buildings and nest boxes.
——— F
1979
Acknowledgments
I thank the co-ordinators of the Canadian nest
records schemes and their co-operators for the use of
data. The nest records are stored at Canadian Wildlife
Service, Sackville, New Brunswick (Maritimes NRS);
National Museum of Natural Science, Ottawa, On-
tario (Quebec NRS); Royal Ontario Museum,
Toronto, Ontario (Ontario NRS); Manitoba Museum
of Man and Nature, Winnipeg, Manitoba (Prairie
NRS); and British Columbia Provincial Museum,
Victoria, British Columbia (British Columbia NRS).
C.S. Robbins read and commented on an earlier
draft, and A. L. A. Middleton and especially M. T.
Myres provided valuable suggestions which aided in
preparation of the final version of this manuscript.
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Boyer, G. F. 1966. Birds of the Nova Scotia~New Bruns-
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ERSKINE: CHANGES IN SWALLOW NESTING SITES
S17
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Received 22 December 1978
Accepted 2 May 1979
Distribution and Habitats of Four Annual Smartweeds in Ontario
RICHARD J. STANIFORTH! and PAUL B. CAVERS2
'Department of Biology, University of Winnipeg, Winnipeg, Manitoba R3B 2E9
2Department of Plant Sciences, University of Western Ontario, London, Ontario N6A 5B7
Staniforth, Richard J. and Paul B. Cavers. 1979. Distribution and habitats of four annual smartweeds in Ontario. Canadian
Field-Naturalist 93(4): 378-385.
Key Words: smartweeds, Polygonum lapathifolium, Polygonum pensylvanicum, Polygonum persicaria, Polygonum
scabrum, distribution, habitats, Ontario.
Polygonum lapathifolium, P. pensylvanicum, P. persicaria, and P. scabrum are closely related and morphologically similar
smartweeds. They often occur together on riverbanks in southern Ontario; outside of riverbanks, their habitats and
distributions differ. Polygonum lapathifolium is found on damp disturbed substrates throughout the province (variety
lapathifolium predominates in the south, and variety salicifolium in the north); P. pensylvanicum is found (as varieties
pensylvanicum and laevigatum) on well drained soils of lakeshores, riverbanks, and occasionally in farmland in southern
Ontario. Polygonum persicaria and P. scabrum have been introduced from Eurasia; in Ontario their distributions are more or
less restricted to wasteland, and cultivated regions where they are locally common as weeds in cereal crops. A key and line
drawings are provided for identification purposes.
Annual smartweeds or persicarias (Genus Poly-
gonum, Section Persicaria) are a group of similar
species which inhabit croplands, as well as wet and
disturbed sites in many parts of the world. In Ontario,
several species are widespread and occur with high
population densities in many locations. The similarity
between species has resulted in their frequent mis-
identification, or in their being lumped together as
“smartweeds.” Furthermore, two or more species may
be found together in close proximity, and this raises
the question of how very similar species can occur
together without one succeeding at the expense of the
others. One possible answer to this question is that
each is predominant in at least one kind of habitat.
The purpose of this study was to clarify and
compare the distribution and habitats of four species
and their varieties in Ontario. The species studied
were Pale Smartweed ( Polygonum lapathifolium L.),
Pennsylvania Smartweed (P. pensylvanicum L.),
Lady’s-thumb (P. persicaria L.), and Green Smart-
“weed (P. scabrum Moench). We follow the nomen-
clature and classification of Scoggan (1978) by recog-
nizing four species and a number of varieties. A
similar classification has been used by Fernald (1950):
however, others (e.g., Gleason and Cronquist 1963)
have preferred to treat P. scabrum Moench as a
variety of P. lapathifolium (P. lapathifolium variety
incanum (Roth) K. Koch). By referring to herbarium
records, we have documented the distribution of these
smartweeds in Ontario. We have also documented,
after an intensive survey, their habitats in Middlesex
County, Ontario.
Methods
Specimens from Ontario were examined from the
following herbaria during 1972 and 1973: OAC,
UWO, TRT, CAN, DAO, SLU, HAM, WLU, WAT,
QK, LKHD, MT, MTJB, MT MG, MICH, MSC, and
the University of Windsor. Morphological data
recorded from these herbarium specimens was later
(1978) used to check specimen identification against
keys in the recently published Flora of Canada
(Scoggan 1978). Each specimen was identified to
variety where possible and its geographical location
was then plotted.
A study area (approximately 3500 ha) was selected
near the village of Delaware, Middlesex County,
Ontario (42°55’N, 81°25’W). A significant criterion in
the selection of this area was that it possessed a variety
of habitats and soil types (Anonymous 1931). The
habitats included cropland (cereals, corn, tobacco,
hay, beans, vegetables), pasture, deciduous wood-
land, roadsides, and river bottomlands. Soil types
included “sands” (Berrien and Fox-fine sandy loams,
Oshtemo and Plainfield sands), “loams” (Guelph,
London, and Parkhill loams and Burford gravelly
loam), and “clays” (Haldimand, Huron, and Perth
clay loams). Sands, loams and clays, as defined above,
occupied approximately equal proportions of the
study area. The Thames River flows southwards
through the center of the area. Many tributaries enter
this portion of the river via gulleys on its east and west
banks.
A survey of the study area was designed to examine
the distribution of the smartweeds in different
378
I)
habitats and on different soil types. The study area
was subdivided into three edaphic types: sand, loam,
and clay; 300 sampling units (each circular with
diameter of 3 m) were placed randomly in each soil
type in such a way that !00 were in cropland, 100 in
woodland, and 100 in disturbed sites other than
cropland (e.g., roadside verges). The presence or
absence of each smartweed species was recorded in
each of the 900 sampling units. If the sampling unit fell
in the wrong habitat, it was moved to the nearest
example of the appropriate habitat. A field check of
the soil type was made at the time that a plot was
placed in the field. In each sampling unit the soil type
was the same as that shown on the Department of
Agriculture soil map (Anonymous 1931).
The frequency of each smartweed species was
determined from 300 sampling units placed along
both sides of a 6.5-km section of the Thames River,
near London, within the study area. Each bank was
divided into 50 sections each 130 10m. Three
circular sampling units, each | m in diameter, were
placed at random within each section. This habitat
was treated separately from those of the previous
survey because riverbank soils could not be classified
as sand, loam, or clay and because the sampling
procedure differed.
Results and Discussion
Distribution in Ontario
Table | gives the numbers of herbarium specimens
STANIFORTH AND CAVERS: ONTARIO SMARTWEEDS
37/9)
examined; most of these were identified to variety
level. Three varieties (P. lapathifolium variety pros-
tratum Wimm., P. pensylvanicum variety durum
Stanford, P. persicaria variety ruderale (Salisb.)
Meisn.) are not recorded by Scoggan (1978) for
Ontario and this is probably the first report of these
taxa for the province. A key (Table 2) and line
drawings (Figure |) are provided for the identification
of Ontario taxa. The key has been modified from keys
in Scoggan (1978) to include only those taxa (to
variety level) found in Ontario, and expanded to
include the previously unreported P. pensylvanicum
variety durum and a key to allow the separation of P.
persicaria variety persicaria from P. persicaria variety
ruderale (Salisb.) Meisn.
Polygonum lapathifolium was the most widespread
species (Figure 2). The most northerly specimen
(CAN 244332) was from Big Trout Lake (53°49’N,
89°53’W). Polygonum lapathifolium has been. col-
lected from all bedrock types, soil types (except
tundra soils), vegetation zones (except tundra), land
use types, and climatic regions of the province.
Variety /apathifolium appears to be the commonest
variety in southern Ontario. Specimens have been
collected from riverbanks, damp agricultural land,
roadsides, ditches, and wasteland. Variety salici-
folium Sibth. is locally distributed in the south, but is
the commonest variety in the northern part of the
province. Specimens of this variety have been col-
lected from sandy beaches of lakes and rivers, and
TABLE |—The varieties of four annual smartweed species found in Ontario, based on herbarium specimens examined by the
authors
No. of specimens
Species Variety examined
Polygonum lapathifolium L. lapathifolium 218
: salicifolium Sibth. 106
prostratum Wimm. eel
325
Polygonum pensylvanicum L. pensylvanicum 81
laevigatum Fern. 46
durum Stanford! 2
eglandulosum Myers? Av
129
Polygonum persicaria L. persicaria 349
ruderale (Salisb). Meisn. as
350
59
Polygonum scabrum Moench
'Specimens appeared to be hybrids between variety durum and variety pensy/vanicum.
2Not encountered in the specimens examined, but reported by Scoggan (1978) for islands in Lake Erie.
380 THE CANADIAN FIELD-NATURALIST Vol. 93
TABLE 2—Key to Polygonum lapathifolium, P. pensylvanicum, P. persicaria, and P. scabrum, and their Ontario varieties. !
This key has been adapted from Scoggan (1978)
1 Ocreae (leaf-sheaths) normally fringed with bristles at summit. Spikes at least 7 mm thick; mature calyx prominently
reticulate at base; achenes prevailingly lenticular; leaves often purplish blotched above; plants mostly of weedy habitats;
(GimMEROG TCE A) ce cree ch etene ss eroe chao ree ates ones ny tue lees ave) oi ceoANfayo shale keane eS PSEIG) OG Casio Lot eR RARE P. persicaria L.
2 Stem ascending or merely decumbent-based; the leaves narrowly to broadly lanceolate; the spike to over
4 cm long var. persicaria
2 Stem prostrate or depressed; the relatively short leaves rhombiclanceolate; the spike usually less than
1.5 cm long var. ruderale (Salisb.) Meisn.
1 Ocreae nearly or quite lacking apical bristles; styles usually 2 (sometimes 3); achenes usually lenticular (sometimes
trigonous)
3 Peduncles and axis of inflorescence with obvious stalked glands or strigose-hispid; spikes pink to purplish (rarely
WANE), Onis? sapAlls Ouseumaly MEA! ocaccosossvenesGesedunooonoOCDOEODLODODDOO EONS P. pensylvanicum L.
4 Peduncles strigose-hispid, without or with very few glands var. durum Stanford?
4 Peduncles and axis of inflorescence covered with gland-tipped hairs.
Si leavesndistinctlyastneoseronmbothisuiblacesm rn ie rice iio ears var. pensylvanicum?
5 Leaves glabrous or at most sparsely strigose on the midrib beneath . var. laevigatum Fern.?
3 Peduncles and axis of inflorescence glabrous or with sessile inconspicuous glands; outer 3 sepals in fruit strongly
3-nerved, each nerve terminating in an anchor-shaped fork (except in P. pensylvanicum var. eglandulosum).
6 Spikes green (rarely purplish), to 5 cm long, erect, the lateral ones mostly sessile or short stalked; floral-axis
copiously glandular; achenes about 3 mm long and about equalling the mature calyx; this not constricted at
the tips(introduced) ei, se iseseisadaitale + se oe eeeoneia tele crs Geeeeysks tees a olsvannere P. scabrum Moench2
6 Spikes pink to purplish.
7 Plant glabrous throughout; calyx pale, about equalling the somewhat shining achene, this is 3-5 mm
longaispikerenectstonl—oremethickeveeemeemiaeeae P. pensylvanicum var. eglandulosum Myers
7 Plant often bearing sessile glands on the peduncles and lower leaf surfaces; calyx pink to purplish,
constricted above into a thick beak overtopping the achene, this is not much over 2 mm long; spike
often somewhat pendulous, not over | cm thick P. lapathifolium L.
8 Leaves lanceolate, broadest near the base, attenuate to tip.
9 Leaves green on both sides, to 2-5 cm long; spikes to 8 cm long, arching or droop-
TTB ses es coe a Sud obese om ae atreri oh en mathe I SU Ree poh oer re var. lapathifolium?
9 Leaves white-pubescent beneath, at most about | cm long; spikes less than 4 cm long:
REC aace ee ean Rea ete ne oe ree var. salicifolium Sibth.?
8 Leaves broadest well above the base, not long attenuate.
Plant prostrate or depressed; leaves subrhombic, mostly not over 7 cm long; spikes to
aboutidicmilom pi cicrenve crstyacle checeher eter mien enna extern var. prostratum Wimm.
'Other annual smartweeds occur in Ontario (e.g., P. caespitosum, P. careyi, P. hydropiper, P. orientale, P. punctatum). These
species are either rare, or more or less restricted to swampy ground (see Scoggan (1978) for treatment of these taxa).
*Specimens of this variety or species which possessed some characteristics of a second variety or species have been seen by the
authors (see text).
from exposed muds. Many intermediates between
these two varieties were noted; for the purposes of
mapping, these were assigned to the variety which
they most closely resembled. A specimen of the variety
prostratum Wimm. (prostrate plant with broad
leaves) had been collected from the vicinity of
Waterloo (WLU 2689).
Polygonum pensylvanicum has the most restricted
distribution of the four species studied (Figure 2). The
most northerly specimen (WAT 718a) was from
Sudbury (45°30’N, 81°00’W). The Ontario distribu-
tion is patchy with three main centers of density:
(a) south of a line from Goderich to Toronto, (6) east
of a line from Kingston to Renfrew, and (c) the
Georgian Bay shoreline. Isolated colonies occur at
Nestorville (46° 18’N, 83°36’W; MICH 5165), North
Bay (46°19’N, 79°28’W; TRT 16735), and Sudbury
(46°30’N, 81°00’W; WAT 718a). Variety pensyl-
1979
POLYGONUM LAPATHIFOLIUM
OCREA PEDUNCLE
SPIKE
STANIFORTH AND CAVERS: ONTARIO SMARTWEEDS 381
3 MM
ACHENE
PERIANTH
FIGURE |. Ocreae, peduncles, spikes, perianths, and achenes (surface and cross sectional views) of annual smartweeds showing
major distinguishing features used in the key (Table 2). Spikes of (a) P. lapathifolium variety lapathifolium, and
(b) P. lapathifolium variety salicifolium. Peduncles of (c) P. pensylvanicum, varieties laevigatum and pensy/vanicum,
(d) P. pensylvanicum variety eglandulosum, and (e) P. pensylvanicum variety hybrid durum X pensylvanicum. Cross
sections of (f) trigonous and (g) lenticular achenes of P. persicaria.
vanicum and variety /aevigatum Fern. are both
common and have similar distributions; however,
their main centers of density differ. Variety pensyl-
vanicum predominates on the Niagara peninsula, but
variety /aevigatum is the most frequent variety in the
vicinity of Ottawa and in Elgin and Middlesex
counties. Many intermediates between these varieties
were noted and, for mapping purposes, were assigned
to the variety which they most closely resembled. Two
specimens were identified by the authors as inter-
mediates between variety durum Stanford and variety
pensylvanicum. These had been collected from Point
Edward, Lambton County (DAO 1862) and Birch
Island, Lake Huron (CAN 44437). No mention is
made of the occurrence of variety durum in Ontario
by Scoggan (1978); however, Gleason (1958) reports a
382 THE CANADIAN FIELD-NATURALIST
POLYGONUM LAPATHIFOLIUM L. var. tapathifolium
POLYGONUM LAPATHIFOLIUM L. var. salicifolium Sibth|
Vol. 93
POLYGONUM SCABRUM Moench
(Pale Smartweed )
POLYGONUM PENSYLVANICUM L. var. pensylvanicum
( Pale Smartweed, willow-leaved variety )
POLYGONUM PENSYLVANICUM L. var. laevigatum Fern
(Green Smartweed )
POLYGONUM PERSICARIA L.
( Pennsylvania Smartweed )
Bi
FIGURE 2. Distribution of four annual smartweeds and their common varieties in Ontario, based on examination of
herbarium specimens.
similar variety intermediate from Indiana. Specimens
of P. pensylvanicum have been collected from
riverbanks, lakeshores, and croplands.
The distribution of P. persicaria was most con-
centrated in the southern half of the province
although specimens have been collected from the clay
plains of Kenora, Dryden, Thunder Bay, Kapuskas-
ing, and New Liskeard. The most northerly specimen
was from Sioux Lookout (50°06’N, 91°55’W). Most
habitats given on specimen sheets were described as
cultivated land or disturbed sites. The species was
recorded from all soil types including Precambrian
deposits. A single specimen of P. persicaria variety
ruderale (Salisb.) Meisn. was identified by the
( Pennsylvania Smartweed, smooth variety)
(Lady's —thumb)
authors. This specimen (WAT 39) had been col-
lected from Conestogo Lake, Waterloo County.
Polygonum scabrum had not been separated from
P. lapathifolium in certain of the herbaria visited.
Data pertaining to peduncle, spike, perianth, and
achene characteristics were used by the authors to
discriminate between these two species using the key
provided by Scoggan (1978). Many specimens pos-
sessed characteristics of both P. scabrum and of P.
lapathifolium (either variety salicifolium or variety
lapathifolium). Such specimens were assigned to the
taxon which they most closely resembled. Polygonum
scabrum is of widespread but local occurrence in
Ontario; it is particularly abundant in the vicinity of
I)
Thunder Bay. The most northerly specimen (TRT
107340) had been collected from Onakawana on the
Moose River. Information on herbarium sheet labels
showed that specimens had been collected from clay
or loam soils of farmland and roadsides, and a few
from riverbanks.
Soil and Habitat Survey
Percentage frequencies for P. lapathifolium and P.
persicaria in each of the nine habitat-soil categories
are shown in Table 3. Polygonum pensylvanicum and
P. scabrum were not encountered in this survey, and
with the exception of one plant of P. persicaria, no
smartweeds were found in woodland.
TABLE 3—Frequency and total number of two smartweeds in
three habitats and on three soil types in Middlesex County,
Ontario. Number of sampling units in each soil type in each
habitat = 100
Species, Brcauencyaa) Total
habitat Sand Loam Clay number
P. lapathifolium
Cropland ] 0 0 |
Disturbed sites 4 2 0 6
Woodland 0 0 0 0
Total number 5 2 0 7
P. persicaria
Cropland 9 24 28 614
Disturbed sites 25 24 24 734
Woodland 1 0 0 ie
Total number 35° 4g>-c Se 135
*4R ow or column totals associated with the same letter are
not significantly different from each other (P > 0.05).
Polygonum lapathifolium was scarce (encountered
in only 7 of 900 sampling units); most occurrences
were in disturbed sites. Frequency values were too low
for statistical comparison. All specimens were of
variety lapathifolium.
Polygonum persicaria was common in croplands
and in disturbed sites on sandy, clay, and loam soils.
The frequency values were sufficiently large to allow
statistical testing of four null hypotheses:
i) That the frequency of P. persicaria on different
soils was independent of its frequency in different
habitats was rejected (P< 0.05) using the G-test
(Sokal and Rohlf 1969). Polygonum persicaria was
most frequent in croplands where these were situated
on clay soils.
ii) That the frequency of P. persicaria did not
differ between soil types, or between cropland and
disturbed sites was tested by means of pairwise
comparisons using the simultaneous testing pro-
cedure (Sokal and Rohlf 1969). The frequency onclay
STANIFORTH AND CAVERS:
ONTARIO SMARTWEEDS 383
soils was significantly higher (P< 0.05) than that on
sandy soils. Frequency values for other pairs of soil
types, and between habitats, cropland, and disturbed
sites were not significantly different.
iii) That the proportion of occurrences of P.
persicaria in cropland increases as one progresses
from sandy through loam to clay soils (i.e., with
decreasing particle size) was accepted (P > 0.05) after
testing of linear proportions (Snedecor and Cochran
1967).
iv) That P. persicaria occurs with equal frequency
in different crop types was tested using an R X C test
of independence based on the G-statistic, and by
pairwise comparisons using the simultaneous testing
procedure (Sokal and Rohlf 1969). These tests took
into account the total numbers of sampling units in
each crop type, in such a way that the scarcity or
abundance of sampling units in a particular crop
species did not affect the results of the test or provide
misleading information. The frequency of P. persi-
caria in small-grained cereals was shown to differ
(P< 0.05) from that in other crops (Table 4). Small-
grain cereals are usually grown onclay or loam soils in
the study area. From these studies it is not possible to
separate the effects of clay and loam soils from the
effects of small-grained cereals on the frequency of P.
persicaria. A further complicating factor is that the
principal crops on drier soils (corn and tobacco) are
usually heavily sprayed with herbicides (such as
Atrazine), to which smartweeds are susceptible.
TABLE 4—Frequency of Polygonum persicaria in various
crop types
Small-grained Other
Crop type Corn cereals! crops?
Number of sites
examined 146 85 69
Number of
occurrences of ic 54> 22
P. persicaria
*’Frequencies associated with the same letter are not
significantly different from each other (P > 0.05).
‘Wheat, barley, oats, rye.
*Tomato, tobacco, beans, melon.
Riverbank Survey
Polygonum lapathifolium variety lapathifolium, P.
pensylvanicum variety laevigatum, and P. persicaria
were encountered in the riverbank survey (Table 5).
The null hypothesis that their frequency values did not
differ was tested by means of a G-test and by
simultaneous testing procedure (Sokal and Rohlf
1969). The null hypothesis was rejected (P< 0.05)
384
TABLE 5—Frequencies of smartweed species along banks of
the Thames River, Middlesex County, Ontario
Species Frequency (%)!
Polygonum lapathifolium 49
Polygonum pensylvanicum 22
Polygonum persicaria 10
Poilygonum scabrum 0
‘Each value is significantly different from each other value
(P< 0.05).
indicating that the values were different from each
other. It is of interest that P. persicaria, the common-
est species in cropland and disturbed sites (habitat and
soil survey), was the rarest of the three riverbank
species. Polygonum pensylvanicum, which was absent
from the previous survey, was common onriverbanks.
Polygonum scabrum was absent from riverbank sites,
as it was from the habitats examined in the previous
survey.
Summary
The distributions and habitats of the four smart-
weeds based on our herbarium studies and field
surveys, are summarized in Table 6.
Polygonum lapathifolium has long been associated
with man’s agricultural activities (Bertsch 1954) and
with man’s assistance has increased its range to most
temperate parts of the world. It is probably not
sufficiently abundant in the croplands of southern
Ontario to be rated as a serious weed in that region. It
is more usually found in wet and naturally disturbed
sites, such as riverbanks, lakeshores, and exposed
mud. It may be abundant in these habitats even at
locations well away from farmland, e.g., in northern
Ontario. When found as a weed, it is usually
associated with damp, disturbed substrates, 1.e.,
conditions very similar to those of the riverbank. Of
THE CANADIAN FIELD-NATURALIST
Vol. 93
the two common Ontario varieties, variety salicifol-
ium is considered to be native but variety /apathi-
folium is believed to be partly introduced (Scoggan
1978). It is not known, however, whether the
introduced element of this variety is the weedy
component.
The range of the native species Polygonum pensyl-
vanicum is more or less restricted to eastern North
America, extending northwards to the Sudbury
district of Ontario (for map, see Staniforth 1975). It is
a serious agricultural weed throughout its range in the
United States, but in Canada it is mostly restricted to
riverbanks and lakeshores. When found as a weed in
Ontario (locally in Kent, Essex, and Elgin counties),
it is associated with crops grown on sandy soils
(especially corn and tobacco). One may speculate that
its northern distribution is correlated with locations
offering a mild climate and light warm soils, such as
those of riverbanks, beaches, and sandy farmland.
Varieties /aevigatum and pensylvanicum are both
common in the province.
Polygonum persicaria, like P. lapathifolium, has
long been associated with man’s agricultural activi-
ties. It has become successfully established in North
America from Eurasia. In Ontario, it is more
restricted to agricultural and man-disturbed habitats
than are the two species discussed above. It does occur
in isolated farming regions such as the clay belt and
the small farmed clay pockets of northern Ontario.
Polygonum persicaria 1s associated particularly with
damp, clay soils in Ontario, especially those which
support crops of small-grained cereals, e.g., oats,
barley, rye, and wheat.
Polygonum scabrum was originally introduced
from Eurasia. It has become widely distributed in
Canada and is now found in all provinces and in the
Northwest Territories (Scoggan 1978). In Ontario, it
is widespread and locally common on damp clay and
loam soils of disturbed substrates, including cropland.
TABLE 6—Summary of the distributions and habitats of four annual smartweeds in Ontario
Distribution,
Species relative abundance
Throughout Ontario;
common
Polygonum lapathifolium
Southern Ontario;
locally common
Polygonum pensylvanicum
Polygonum persicaria
province; common
Polygonum scabrum
All cultivated parts of the
Throughout Ontario; local
Principal
habitats
Principal
soil type
Riverbanks, lakeshores,
sometimes farmland and
waste places
Damp clays and loams
Riverbanks, lakeshores,
locally in crops
Well-drained loams,
sands, and gravels
Wasteland, cultivated land
(cereal crops), occasionally
riverbanks
Damp clays and loams
Wasteland, cultivated land,
occasionally riverbanks
Damp clays and loams
ee
1979
It was not encountered during habitat surveys in
Middlesex County, Ontario.
Acknowledgments
We thank the Canadian Northern Sportsmen’s
Show for financial support of this project. Comments
offered by the referees were appreciated. The assis-
tance and co-operation of curators of herbaria in
Ontario, Quebec, and Michigan are also gratefully
acknowledged. In addition, we thank H. Scoggan for
permission to use his keys to species and varieties of
Polygonum.
Literature Cited
Anonymous. 1931. Soil survey map of the County of
Middlesex, Province of Ontario. Soil Survey Report
Number 6, Experimental Farm Services, Department of
Agriculture, Ottawa.
Bertsch, K. 1954. Vom neolithischen Feldbau auf der
Schwabischen Alb. Berichte der Deutschen Botanischen
Gesellschaft 67: 18-21.
Fernald, M. L. 1950. Gray’s manual of botany. 8th Edition.
STANIFORTH AND CAVERS: ONTARIO SMARTWEEDS
385
American Book Company, New York. 1632 pp.
Gleason, H. A. 1958. The new Britton and Brown illustrat-
ed flora of the northeastern United States and adjacent
Canada. Volume 2. Revised edition. New York Botanical
Garden. 655 pp.
Gleason, H. A. and A. Cronquist. 1963. Manual of vascu-
lar plants of the northeastern United States and adjacent
Canada. Van Nostrand Company, Princeton. 810 pp.
Scoggan, H. J. 1978. Flora of Canada. Part 3. National
Museums of Canada, Publications in Botany Number
7 (3). 568 pp.
Snedecor, G.W. and W.G. Cochran. 1967. Statistical
methods. 6th Edition. Iowa University Press, Ames.
593 pp.
Sokal, R. R. and F. J. Rolf. 1969. Biometry. The principles
and practice of statistics in biological research. W. H.
Freeman and Company, San Francisco. 776 pp.
Staniforth, R. J. 1975. The comparative ecology of three
riverbank annual plants. Ph.D. thesis, University of
Western Ontario, London, Ontario. 368 pp.
Received 6 September 1978
Accepted 29 May 1979
Apparent Differences in Aquatic Macrophyte Floras of Eight
Lakes in Muskoka District, Ontario from 1953 to 1977
G. E. MILLER and H. M. DALE
Department of Botany and Genetics, University of Guelph, Guelph, Ontario NIG 2W1
Miller. G. E.and H. M. Dale. 1979. Apparent differences in aquatic macrophyte floras of eight lakes in Muskoka District.
Ontario from 1953 to 1977. Canadian Field-Naturalist 93(4): 386-390.
Forty-eight species of aquatic macrophytes were recorded in two surveys made 23 yr apart of the same areas in eight
Muskoka, Ontario lakes. Only 69% of 265 sightings were duplicates in the surveys. Improved sampling methods resulted in
additional records of deep-water species whereas certain floating-leafed species were no longer extant in some lakes. Some
currently obvious species are thought to have changed sufficiently in abundance to produce the new records. In the shallow-
water records. the changes in several subterranean and inconspicuous species were attributed to presence or absence of
obvious flowers.
Key Words: aquatic plants, macrophyte, flora, biological surveys. freshwater lakes, long-term changes. Muskoka District,
Ontario.
The process of eutrophication brings about changes
in the aquatic flora in a short time (Lind and Cottam
1969: Harman and Doane 1970). Undisturbed aquatic
stands, rephotographed in Scotland by Spence (1964)
to compare with earlier photographs by West (1905)
showed no significant successional change in aquatic
communities after more than 50 yr. Dramatic changes
more frequently result from man’s activities, because
they may alter the water chemistry, clarity, and
temperature: these factors have been linked with
species change (Dale and Miller 1978: Stuckey 1971:
Volker and Smith 1965).
The scarcity of accurate floristic surveys of lakes
hinders the weighing of the reality of an apparent
change in a flora. Too often published records are
based on traditional techniques using remote sam-
pling from the water surface, with a rake or a
grappling hook (Shields!: Soper: Stuckey 1971), or
an Ekman dredge (Rich et al. 1971). Direct examina-
tion of aquatic vegetation (ina manner comparable to
those in which observations are made in terrestrial
ecology) using SCUBA was introduced by Schmid
(1965) and was the method used for this study.
The eight lakes studied are of the unbuffered,
slightly acid type, characteristic of the Precambrian
Shield of central Ontario. Three Mile Lake has an
alkalinity of 9 mg CaCO,/L and an electrical con-
'Shields, J.K. 1953. A survey of the aquatic and marsh vege-
tation of some lakes in the Muskoka and Parry Sound
Districts of Ontario. A report prepared for the Toronto
Fish and Game Protective Association, Toronto.
"Soper, J.H. 1948. A survey of the aquatic vegetation of
Whitewater Lake, with special reference to its suitability to
waterfowl. A report prepared for the Department of Lands
and Forests of Ontario, Toronto. 13 pp. mimeo.
ductivity of 56 wzmho/cm. The other seven lakes have
alkalinities below 5 mg CaCO;/L and conductivities
in the range of 29-41 wmho/cm. All pH readings were
in the range 6.0-6.6. Three Mile Lake has a notice-
ably larger clay component in its substrate; reduced
water clarity resulted in a 2.1-m Secchi disc reading
compared with the 3.9—9.4 m in the other lakes. Lakes
such as these characteristically have both a low
productivity of aquatic macrophytes and a low
standing crop when compared with lakes richer in
dissolved solids. It 1s not known whether these values
have changed between the two sampling dates.
Differences in the recorded flora are assessed as
attributable to sampling technique, to man’s influence
on the environment, or to successional trends in the
vegetation.
Methods
Three two-day surveys (by motor boat) were made
of each lake by a team of three in the summer of 1976.
Several stands (10 to 20) were carefully surveyed using
SCUBA or skin-diving equipment. Sampling sites
were chosen so that areas fully described in the former
survey on the maps of Shields! were revisited. In
addition, sites of unique features as well as ones
characteristic of the lake were included. The vegeta-
tion in an area 1000 m2? was surveyed and all
submersed and floating aquatic plant species were
rated for abundance on the scale: 0, absent to 4,
abundant. Each species was given a mean abundance
rating for each lake using the same scale. This index
indicated the frequency of an expected encounter. The
herbarium at the University of Guelph (OAC) houses
a sample of each species. Five of the eight lakes were
revisited in 1977 to check typical locations for species
apparently ‘lost’ since 1953.
386
1979 MILLER AND DALE: MACROPHYTE FLORAS OF MUSKOKA LAKES 387
80°W 79°'W
(1)
\ NORTH BAY
LAKE NIPISSING
BEATTY LAKE ro)
46°N t)
4 Trout Creek 46°N
EAGLE L
HORN LAKE
D
Burk’s Falls
RAINY LAKE P
BUCK LAKE
PARRY SouND 4%. 44KE
SKELETON
ey) LAKE
THREE
MILE
LAKE
GEORGIAN BAY
45°N 45° N
Gravenhurst
80°W 79°W
FIGURE |. Location of the eight lakes sampled. Alternate names for some of these lakes are Rainy (Bartlett). Horn (Sollman).
and Eagle (Machar).
388
Results
Forty-seven species were identified; this is 12 more
than in 1953! (Table 1). There was a loss of a single
species of Pondweed (Potamogeton obtusifolius).
Five species present in the total flora were missing
from two or more lakes; of these, four produce floating
leaves and the fifth, a bladderwort, is chiefly subter-
ranean but becomes noticeable when it produces
yellow aerial flowers.
Discussion
Although the methods of sampling and recording
the data differed, the two studies are comparable. The
thorough survey, using a rowboat and rake to
determine species present and using symbols to record
on maps, produced a complete sampling for the parts
of the lake visible from the surface. The recent survey
used the same maps to check stretches of shallow
water barren of plants in 1953 as well as areas in which
a complex of species was found. These lakes have long
stretches with few plants and the richest communities
consisted of less than a dozen submersed and floating
species. The numerous sites were surveyed by wading
and swimming by a team of three but there is no
reason to suggest that this was an incomplete survey of
the shallow-water communities. The major difference
in the surveys was in a more complete sampling of the
deeper water using SCUBA equipment.
Of the plant species that were missing from two or
more lakes, Batchelder (G/yceria borealis), a plant of
small scattered patches, is difficult to identify after the
release of seed, as is the bladderwort (Utricularia
cornuta) if it is not in flower. One site of the latter was
lost when a highway was built over a small part of
Eagle Lake. Three others, the floating-leafed species,
must be lost from the lake floras. These very visible
species could not be overlooked and their loss was due
to severe reduction of their populations, perhaps
because of the increased use of outboard boat motors.
The effect of the propellors would be similar to that of
a mowing machine constantly cutting away the leaves.
The number of buildings on one stretch of shore in
Skeleton Lake increased from 102 1n 1956-1960 to 177
in 1970 (Map 31 E/3 west. Ist edition, 1960 and
new edition, 1974. National Topographical System,
Canada). The decades between surveys were also the
years of great water skiing activity in the Muskoka
lakes.
Plants recorded only in the second survey were
considered indicators of succession if found in two or
more lakes. There has been no study to show whether
Quillwort (/soetes macrospora), Big-leaf Pondweed
(Potamogeton amplifolius), and Arrowhead (Sagit-
taria graminea) could have increased dramatically in
abundance in the interval. It is probable that these
THE CANADIAN FIELD-NATURALIST
Vol. 93
species were present but only in deep water and were
not reached by the rake in the first sampling. For the
same reason, six species — Quillwort, Arrowhead,
Water moss ( Fontinalis sp.), Bushy Pondweed ( Najas
flexilis), Big-leaf Pondweed, and Purple Bladderwort
(Utricularia purpurea) — could also have been un-
recorded. The last species was unrecorded in Law-
rence Lake from a surface-based sampling (Rich et al.
1971) although it was common in part of the lake in
1976 (H. M. Dale and D. H. N. Spence, unpublished
data).
Two of the new species, the bladderworts U.
resupinata and intermedia, are obscure and could
have been overlooked in 1953. The remaining species,
however, are widespread shallow-water species, read-
ily seen and identified. These species, therefore, have
become more conspicuous or are new elements in the
flora.
Along the southern shore of Three Mile Lake there
is a horseshoe-shaped bay described by Shields! on his
maps as containing a dozen species including six with
floating leaves. This location was found to be
unchanged in 1976 and matched the earlier descrip-
tion exactly. In contrast, a site along the western shore
of Eagle Lake was marked clearly in 1953 as occupied
by only Seven-angled Pipewort (Eriocaulon sept-
angular) and Water Lobelia (Lobelia dortmanna), yet
in 1976, a very conspicuous population of Narrow
Floating-leaf Bur Reed (Sparganium angustifolium)
was growing from a dense mat of the former two
species. Also present were Needle Rush (Eleocharis
acicularis), Rush (Juncus pelocarpus), and Waterwort
(Elatine minima), members of the promiment wide-
spread new species. Similarly in Horn Lake, Floating-
heart and Floating-leaf Bur Reed had invaded a
Pipewort-Lobelia mat.
In this type of sandy oligotrophic lake, Seven-
angled Pipewort and Water Lobelia act as colonizing
species and form patches that stabilize the coarse
substrate, trap fine particles, and increase the organic
matter of the substrate. These patches grow and
coalesce, forming a modified environment suitable for
colonization by Bur Reed. Waterworts, Rush, Needle
Rush, and Floating-heart. Such a successional pro-
cess in the vegetation produces a more complex
vegetation by increasing the abundance of the rarer
species.
Some differences between the surveys 23 yr apart
may be explained by improved sampling methods:
sampling with the use of a rake is less accurate than
using SCUBA equipment. The floating-leafed Yellow
Water Lily, Water Shield, and Floating-heart have
been lost as a result of recreational use of the water:
other species increased in abundance as a result of
succession.
1979 MILLER AND DALE: MACROPHYTE FLORAS OF MUSKOKA LAKES 389
TABLE |—Prescence (+) or abundance (R—rare, O—occasional, C—common, A—abundant) of aquatic macrophytes at the
two sampling dates (1953, 1976) in each of eight lakes. Common names are from Fassett (1957)
Three Skeleton Axe Buck Rainy Horn Eagle Beatty
Mile Lake Lake Lake Lake Lake Lake Lake
Aquatic macrophytes os Ho 33 WO s3 oO 33 Oo 933-10 ~ 33 10 33 1 38 16
Water Moss (Fontinalis sp.) cnn Rec erect tome FY O (S O
Quillwort (/soetes macrospora) O C C A Cc A +
Narrow Floating-leaf BurReed (Sparganium
angustifolium) R C +
Floating-leaf BurReed (Sparganium fluctuans) + R + O OF
Big-leaf Pondweed (Potamogeton
amplifolius A
Narrow-leafed Pondweed ( Potamogeton
berchtoldii)
Ribbon-leaf Pondweed (Potamogeton
epihydrus) ap JR ae
Variable Pondweed ( Potamogeton
gramineus) +
Floating Brownleaf (Potamogeton natans) tone Ati:
Small Floating-leaf Pondweed ( Potamogeton
oakesianus)
Pondweed (Potamogeton obtusifolius) +
Sago Pondweed (Potamogeton pectinatus) R
Clasping-leaf Pondweed ( Potamogeton
richardsonii) +
Robbins’ Pondweed ( Potamogeton robbinsii)
Pondweed (Potamogeton spirillus)
Bushy Pondweed (Najas flexilis)
Arrowhead rosettes (Sagittaria graminea) +
Canada Waterweed (Elodea canadensis)
Wild Celery (Vallisneria americana)
Batchelder (Gl/yceria borealis)
Wild Rice (Zizania aquatica)
Needle Rush (Eleocharis acicularis) A Cc R
Triangle Spike Rush (Eleocharis robbinsii)
Water Bulrush (Scirpus subterminalis)
Water Arum (Calla palustris)
Seven-angled Pipewort (Eriocaulon
septangulare) romeo O) bmi ttt ane Cpa
Pickerel Weed (Pontederia cordata)
Bayonet Rush (Juncus militaris)
Rush (Juncus pelocarpus)
Smartweed (Polygonum amphibium) +
Coontail (Ceratophyllum demersum)
Water-shield (Brasenia schreberi) +
Yellow Water Lily (Nuphar variegatum)
White Water Lily (Nymphaea odorata)
Creeping Spearwort (Ranunculus reptans)
Water Starwort (Callitriche heterophylla) 1
Waterwort (Elatine minima)
Mares-tail (Hippuris vulgaris) +
Farwell’s Milfoil (Myriophyllum farwelli)
Milfoil (Myriophyllum tenellum) +
Floating-heart (Nymphoides cordatum) +
Bladderwort (Utricularia cornuta)
Bladderwort (Utricularia gibba)
Bladderwort (Utricularia intermedia)
Purple Bladderwort (Utricularia purpurea)
Bladderwort ( Utricularia resupinata)
Common Bladderwort (Utricularia vulgaris) an Reet
Water Lobelia (Lobelia dortmanna) + erie Cathe ae 3
Number of changes/ total records 6/22 7/20 10/1
QO 7 non OS)
+
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+
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+
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+
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+
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+
+
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+ ++ +
ARN S
+
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a
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+
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+
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+
+
Aw
+
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+
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+
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+
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i
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+
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+ + + + +
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+
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12/25 18/32 9/17 l
oS)
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oo
~
390
Acknowledgments
We are grateful to R. Dreimanis and B. Elder for
their cheerful assistance during the initial sampling.
This study was partially supported by National
Research Council of Canada grant 3402.
Literature Cited
Dale, H. M. and G. E. Miller. 1978. Changes in the aquatic
macrophyte flora of Whitewater Lake near Sudbury,
Ontario from 1947 to 1977. Canadian Field-Naturalist 92:
264-270.
Fassett, N.C. 1957. A manual of aquatic plants. 2nd
edition, with revision appendix by E. C. Ogden. Uni-
versity of Wisconsin Press, Madison, Wisconsin.
Harman, W.N. and T.R. Doane. 1970. Changes in the
aquatic flora of Otsego Lake between 1935-1939. New
York Fish and Game Journal 17: 121-213.
Lind, C. T. and G. Cottam. 1969. The submerged aquatics
of University Bay: A study in eutrophication. American
Midland Naturalist 81: 359-369.
Rich, P.H., R.G. Wetzel, and N. Van Thuy. 1971.
Distribution, production and role of aquatic macrophytes
THE CANADIAN FIELD-NATURALIST
Vol. 93
in a southern Michigan marl lake. Freshwater Biology
1: 3-21.
Schmid, W. D. 1965. Distribution of aquatic vegetation as
measured by line intercept with SCUBA. Ecology 46:
816-823.
Spence, D.H.N. 1964. The macrophytic vegetation of
freshwater lochs, swamps and associated fens. Jn The
vegetation of Scotland. Edited by J. H. Burnett. Oliver
and Boyd, Edinburgh and London. pp. 306-425.
Stuckey, R. L. 1971. Changes of vascular aquatic flowering
plants during 10 years in Put-in-bay Harbour, Lake Erie.
Ohio. Ohio Journal of Science 71: 321-342.
Volker, R. and S. G. Smith. 1965. Changes in the aquatic
vascular flora of Lake East Okoboji in historic times.
Proceedings of the Iowa Academy of Science 72: 65-
12.
West, G. 1905. A comparative study of the dominant
phanerogamic and higher cryptogamic flora of aquatic
habit in Scottish lakes. Proceedings of the Royal Society
of Edinburgh 25: 967-1023.
Received 17 January 1979
Accepted 28 March 1979
Utricularia geminiscapa at Mer Bleue and Range Extensions in Eastern
Canada*
ERICH HABER
Botany Division, National Museum of Natural Sciences, National Museums of Canada, Ottawa, Ontario KIA 0M8
Haber, Erich. 1979. Utricularia geminiscapa at Mer Bleue and range extensions in eastern Canada. Canadian Field-
Naturalist 93(4): 391-398.
The first Ontario occurrence of Utricularia geminiscapa at the Mer Bleue peat bog just east of Ottawa is reported. The main
features that help to distinguish this species from U. vulgaris are tabulated and the problem of differentiating vegetative
specimens of these two species is discussed. Additional range extensions for U. geminiscapa in eastern Canada of up to 600 km
north of previously known localities are also reported and plotted on a distribution map of the species for North America.
Key Words: Utricularia geminiscapa, morphology, range extensions, Ontario, Canada.
The discovery of an Ontario locality for the Research Council fellow with the National Museum
bladderwort Utricularia geminiscapa occurred onone _ of Natural Sciences, and I were returning to our access
of several excursions made in 1973 to Mer Bleue, a __ point at the tip of Dolman Ridge when hé noticed
peat bog situated approximately 16km east of some flowering Utricularia in a small bog pool.
Parliament Hill, Ottawa (Figure 1). David R. Given Because it was obviously different from the U.
from New Zealand, who was then a National intermedia we had seen earlier, we made a repre-
FIGURE 1. Mer Bleue peat bog, Ontario. Dolman Ridge is the northern spit of land extending into the bog from the left,
Borthwick Ridge the southern.
*Contribution Number 9 to “Scientific and Cultural Studies of the Mer Bleue.”
39
392
sentative collection. The identity of this species was
readily determined at a later date because it had the
characteristic chasmogamous and_ cleistogamous
flowers (Figure 2a). Indeed the seeds from the
cleistogamous flower capsules (Figure 2b) proved to
be extremely characteristic and easily referable to U.
geminiscapa by comparison with seed illustrations in
Muenscher (1944). This initial encounter with U.
geminiscapa, a species new to the province, prompted
me to examine the morphology and habitat character-
istics of this bladderwort in more detail and to search
for additional range extensions in eastern Canada.
Morphological Distinctions
Although U. geminiscapa is usually quite distinct
from U. vulgaris, particularly when flowers or
capsules are present, purely vegetative specimens of
these species may at times be difficult to identify
unequivocably. The vegetative plumes of foliage of
small specimens of U. vulgaris can approach in width
those of U. geminiscapa, which are usually much
narrower than those of U. vulgaris. The identification
of vegetative material is further complicated by the
lack of a clear-cut distinction in the form and nature of
the leaf branching in the two species. In U. gemi-
niscapa, the branching is dichotomous (Figure 3a)
and is usually distinct from the pseudopinnate
branching found in normal robust specimens of U.
tT eee “rf eee
2 aN — |
4 va . é vg ’ \
ae
e \
Sie 2 Sem
THE CANADIAN FIELD-NATURALIST
Vol. 93
vulgaris (Figure 3d). This distinction, however, is not
always evident because smaller specimens of U.
vulgaris may exhibit leaf branching similar to that of
U. geminiscapa. Such vegetative characteristics are
often almost obliterated by careless mounting pro-
cedures in the preparation of herbarium specimens.
Fresh specimens should be float-mounted prior to
drying to display clearly the nature of the leaf
branching. Several features that can be used to
distinguish these two species are given in Table 1.
The supposed absence of marginal leaf spines in U.
geminiscapa has been used as a criterion for distin-
guishing this species from U. vulgaris which is
characteristically spinulose-margined (Muenscher
1944: Fassett 1957: Roland and Smith 1969). The
fresh specimens from the Mer Bleue locality leave no
doubt that the leaf segments can be spinulose-margined
(Figure 3a). The abundant herbarium specimens from
the Farnham bog (Missisquoi County, Quebec) also
confirm this observation. The leaves of the winter buds
are also slightly spinulose and are terete in section, as in
normal leaves. They forma loose, one-sided aggregate
and maintain their green color when in bud (Figure
3b). In U. vulgaris the bud leaves are usually
conspicuously flattened, highly dichotomized and
very spinulose with the whole bud appearing grayish-
white owing to the abundance of the solitary or
fasciculate spines. The bud forms a dense, round to
a
-
&
\ —
\ oe
\ Pe
t et
eA # Y
ae = 3 © x : ty
. 2 * te es
tte : ia,
; Y a _ x
x SO . - * * oe 12
7 3 aes a yes
‘ Lae & = ys
‘A ‘\ :
ov h :
= 2
wt
: Pe
4 * Oy
< 9
hve a
ees Y
a
FIGURE 2. Utricularia geminiscapa: (a) whole plant with normal flowers and cleistogamous capsules, the latter indicated by
arrows; (b) seeds from cleistogamous capsules.
1979 - HABER: UTRICULARIA GEMINISCAPA IN EASTERN CANADA 393
FIGURE 3. Leaf branching pattern and winter bud characteristics in Utricularia. Utricularia geminiscapa: (a) spiny leaf
segments in a fresh specimen; (b) portion of leafy plume of a fresh specimen with terminal winter bud (the one-sided
aspect of the bud is not evident in this photo). Utricularia vulgaris. (c) portion of leafy plume of a dried specimen with
terminal, highly spinulose winter bud; (d) leaf branching pattern in a dried specimen.
394
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 1—Characters useful in distinguishing the two species of Utricularia
Character
filiform, without scales
corolla 0.5-0.8 cm long
without basal lobes
Aerial scapes
Flowers
Flower bracts
Cleistogamous flowers
U. geminiscapa
present, apetalous, scattered along stem
U. vulgaris
coarse, with several scales
corolla 1.5-2.5 cm long
with basal lobes
absent
axils and producing abundant seeds
discoid
1.0-3.5 cm in diameter
dichotomous
Seeds
Foliage plumes
Leaf branching
Leaf segments
spinulose
Winter buds
ingly spinulose
long acuminate with or without terminal
spine(s); margins glabrous to sparingly
0.2-0.5 cm in diameter, green, one-sided;
leaf segments terete, glabrous to spar-
angular with many facets
3-12 cm in diameter
dichotomous to usually + pinnate
spine-tipped with single or fasciculate
spines; margins + abundantly spinulose
1-3 cm long; dense, round to oval mass,
occasionally lobed; leaf segments; flat-
tened and highly spinulose
oval, sometimes lobed mass of bud leaves (Figure 3c).
The best way of ensuring that vegetative specimens of
these two species can be identified with some certainty
is to collect from a given locality samples including as
wide a size variation of the vegetative plumes as
possible.
Although U. intermedia also occurs in bog pools at
Mer Bleue, it can be readily distinguished from U.
geminiscapa and as well from U. vulgaris by its
flattened, dichotomous branching and by the presence
of a mid-vein in the leaf segments.
Characteristics of the Mer Bleue Locality
A search of herbaria in Ontario and Quebec
indicated that the Mer Bleue site is probably the first
verified locality for U. geminiscapa in Ontario (see
also Morris 1922). In order to gain an insight into its
frequency of occurrence at the bog, I returned during
October and early November of 1978. The water level
in the fall of 1978 was somewhat lower than when the
first collection was made (17 July 1973) with the result
that the original shallow pool was no longer present.
In its place were several small potholes with little
standing water and no discernible evidence of the
bladderwort. The yearly and seasonal water table
fluctuations, therefore, appear to have some influence
on the localized development of U. geminiscapa in
shallow bog pools. Presumably low water levels
would not eradicate the species from such shallow,
superficial bog pools because of the presence of the
resistant winter buds which could lie dormant in the
moist Sphagnum mat until open water was again
available.
Potential sites for the bladderwort could be readily
located by inspecting aerial photographs of Mer
Bleue. The pools of varying size that are evident on the
aerial photographs are distributed primarily in the
vicinity of the wooded islands in the central region of
the bog (see Figure 4). Most if not all of these are
bomb craters formed when the bog was used as a
practice bombing range between 1942 and 1945 (S.
Ashley, 1979. The Mer Bleue — The evolution of an
urban bog. An oral history of the Mer Bleue and
surrounding communities. An in-house publication,
National Capital Commission, Ottawa, Ontario;
Freeman 1969). The latest aerial photographs taken in
1978 (Figure 4 in part) indicate that virtually all of the
pools evident on the earliest photographs taken in
1945 are still present. x
Five of the six bomb crater pools surveyed briefly in
the fall of 1978 contained the bladderwort (sites 6, 9,
12, 13, 14; Figure 4). One of these pools (site 14)
located in a predominantly open area with scattered
Tamarack (Larix laricina) is illustrated in Figure 5.
These deep, bomb crater pools contain few aquatics
and are usually fringed by a relatively firm Sphagnum
mat stabilized by ericoid shrubs, in particular by
Leatherleaf (Chamaedaphne calyculata). The pres-
ence of U. geminiscapa in nearly every pool sampled
would indicate that the unorthodox use of the bog asa
bombing range may have benefited the species by
increasing the number of habitats suitable for coloni-
zation. These deep pools would also serve as favored
habitats for this bladderwort because low water-table
levels would not eliminate such pools.
Water samples were collected from each pool
surveyed and the pH compared with bog water
obtained from within the bog mat at various sites. The
location of these samples is indicated in Figure 4.
Summarized in Table 2 are the pH measurements
which were obtained in the laboratory at room
temperature with a Beckman 180 pocket pH meter.
The pH values obtained ranged from 3.9 to 6.1. Sites
1-5 and 7 with pH values ranging from 3.9-4.] are
perhaps most representative for the bog mat, judging
from reports by Terasmae and Mott (1964, pH 3.8)
1979 HABER: UTRICULARIA GEMINISCAPA IN EASTERN CANADA 395
FIGURE 4. Original collection locality for Utricularia geminiscapa (asterisk). It was subsequently found at sites 6, 9, 12, 13, 14.
Each numbered site represents a sample locality for pH determinations.
FIGURE 5. Northward view of one of the deep, bomb crater pools (site 14) in which Utricularia geminiscapa was found.
Photographed in the fall of 1978.
396
TABLE 2—Acidity (pH values) of pools and bog mat at
various sites in the Mer Bleue peat bog
Site pH
I open mat 4.1
2 open mat 4.0
3 forested bog 39)
4 forested bog 38)
5 sedge zone 4.0
6 bog pool 4.5
7 sedge meadow 4.1
8 bog pool 6.1
9 bog pool 5.4
10 open mat 4.7
11 sedge-leatherleaf mat 4.6
12 bog pool 49
13 bog pool 4.9
14 bog pool 47
15 sedge-leatherleaf mat 4.4
16 marginal channel (lagg) 4.6
and Joyal (1971, pH 3.8 and 3.9). The bog pools
ranging from 4.5 to 6.1 had consistently higher values
than sites in the bog mat. The remaining sites (10, 11,
15, 16) also had higher values (4.44.7) than the more
usual value of about 3.9 for the bog mat. Measure-
ments previously taken in June 1974 in the region east
of site 16 and extending to the first small island yielded
pH values between 5.1 and 5.4.
The significance of the higher pH levels of the bomb
crater pools and other sites in the central region of the
bog is not clear until one also takes into account the
localized enrichment of the bog flora that occurs in
this area. Some of the species that occur here are also
common elements of the peripheral eutrophic zone.
This central, primarily treeless, mesotrophic region is
represented in Figure 4 as the dark elongate band
extending eastward from the southeastern tip of
Dolman Ridge to the first small island and again as
the major band surrounding and trailing eastward
from the large central islands. Cat-tails (Typha
latifolia), Bogbean (Menyanthes trifoliata), Water
Arum (Calla palustris), Chain Fern (Woodwardia
virginica), and Scheuchzeria palustris occur 1n this
mesotrophic region. The White Fringed Orchid
(Platanthera blephariglottis) is also common along
the southern, forested edge of this area east of the
large islands (see also Lafontaine 1971). Speckled
Alder (Alnus rugosa), a common bog fringe species,
occupies a sizable shrubby area between the western
ends of the two large islands, and species of sedges
form extensive meadows in this central region and add
to the fen-like appearance.
The generally higher pH values and apparent
mesotrophic nature of this central region may be
THE CANADIAN FIELD-NATURALIST
Vol. 93
directly related to the ground-water levels and shallow
nature of the underlying central basin (maximum
depth about 2 m) (see map of peat depths in Nystrom
and Anrep 1909). During periods of drought as in
1877 and 1878 (The Ottawa Naturalist 1888, 2: 74) and
low ground-water levels as in 1970 and 1974 (M. Bik,
1975. In Proceedings of the Mer Bleue Seminar. An
in-house publication, National Capital Commission
and Central Research Forest, Ottawa, Ontario), the
water level would decrease over the central shallow
basin and aeration and substrate temperatures would
increase resulting in a temporary eutrophication.
Subsequent periods of high ground-water levels
would cause a reversai of these conditions. This
mesotrophic zone lying between two deeper basins
(maximum depths about 6 m) may in effect represent
a tension zone providing suitable habitats for certain
species of plants. The relatively large band of Typha
within this zone east of the large islands may
represent, together with the peripheral 7ypha zone,
the remnants of an earlier, more extensive marshland
habitat. In Camfield’s pollen diagram (1969), Typha
pollen was most abundant in one of the oldest zones in
her Mer Bleue core.
Although the earliest organic sediments in Mer
Bleue are approximately 7500 yr old (Camfield 1969),
the arrival of U. geminiscapa probably dates to a
much more recent time commensurate with the
formation of suitably acidic bog pools. Throughout
its range this species occurs most commonly in quiet,
acidic waters and in particular in bog pools. The
difference in habitat preferences between U. gemi-
niscapa and U. vulgaris is readily apparent at Mer
Bleue where the latter occurs in shallow ditches
adjacent to the bog but not directly influenced by the
acid Sphagnum mat. Throughout its widespread
range, U. vulgaris occurs in ponds, lakesides, and
sluggish streams.
The occurrence of U. geminiscapa at Mer Bleue is
simply one more interesting species addition to an
increasing list of novelties either first described from
the bog or represented regionally only at this locality.
Additional information regarding various aspects of
this locality can be found in the papers by Baldwin and
Mosquin (1969), Corbet and Walley (1969), Dunston
(1970), Smith (1970a, b), Hobson(1970), Ouellet et al.
(1976), and Small (1976).
Range Extensions in Eastern Canada
A search of herbaria in Ontario and Quebec (CAN,
DAO,. TRT, QFA, QSA, MT, MIMG, and SEs:
acronyms according to Holmgren and Keuken (1974))
resulted in the validation of specimens from seven
Canadian localities in addition to the Mer Bleue
record. The eight validated localities of U. gemi-
niscapa plotted on the map in Figure 6 are as follows:
FIGURE 6. North American distribution of Utricularia
geminiscapa. The generalized range of this species,
derived from maps published by Fernald (1933),
Thomson (1940), Muenscher (1944), and Roland and
Smith (1969), is represented by the hatched areas. The
solid dots represent new localities based on verified
specimens.
Ontario: Mer Bleue peat bog, 45°24’N, 75°30’W,
The Regional Municipality of Ottawa-Carleton, bog
pool, E. Haber & D. R. Given 1963, 17 July 1973
(CAN).
Quebec: plaine de la Riviere Ouelle, cté Kamouraska,
E. Campagna, 13 July 1928 (QSA); tourbiére de
Farnham, cté Missisquoi, M. Raymond, 20 Aug. 1933
(CAN) [the earliest of many collections from this
site]; L. Hibou, 5 mi[3 km] southwest of Taschereau,
W. Abitibi Co., small boggy lake, W. K. W. Baldwin
& A. J. Breitung 4287, 29 Aug. 1952 (CAN); entre St-
Faustin et Lac Supérieur, Lac artificiel Dupré, cté
Terrebone, Frére Rolland-Germain 422, 23 July 1957
(CAN); Ilets-Jérémie, cté Saguenay, dans les petites
mares, tourbiére a cypéracéés, J. Cayouette 73-719,
14 Aug. 1973 (QFA); Hopkins Hole, between Ramsay
L. and Hawley L., Gatineau Pk., Gatineau Co., A. &
J. Reddoch, 19 Aug. 1978 (CAN).
The most northerly range extensions for U. gemi-
HABER: UTRICULARIA GEMINISCAPA IN EASTERN CANADA
397
niscapa occur at Lake Hibou (48°38’N, 78°48’W) and
at Ilets-Jérémie (48°53’N, 68°48’W) jn Quebec. Both
localities are approximately 600 km north of pre-
viously documented localities in upper New York
State and Maine, respectively. Although the Lake
Hibou specimens were originally determined as U.
minor by Baldwin, they are sufficiently characteristic
in spite of their vegetative condition that they were
validated as U. geminiscapa.
A number of perplexing collections made by Frére
Rolland-Germain at Lake Monroe in Montcalm
County, Quebec and identified as U. geminiscapa
(CAN, MT, SFS) are considered to be merely small
vegetative specimens of U. vulgaris. This decision was
reached only after seeing a whole series of vegetative
specimens of U. vulgaris from the same locality at the
Marie-Victorin herbarium (MT). These specimens
ranged from the more typical large-plumed specimens
to those that were much reduced in size approaching
U. geminiscapa in appearance.
Utricularia geminiscapa may indeed be much more
widely spread in eastern North America than is at
present documented. The vegetative characteristics of
U. vulgaris such as overall size of leaf plumes, manner
of leaf branching, form and pubescence of winter bud
leaves, and leaf pubescence are extremely variable and
require further investigation over the whole range of
this cosmopolitan species. The cultivation of this
species under controlled environmental conditions
may prove useful in accounting for some of the
observed natural variation.
Literature Cited
Baldwin, W. K. W. and T. Mosquin. 1969. Scientific and
cultural studies of the Mer Bleue. Canadian Field-
Naturalist 83(1): 4-6.
Camfield, M. 1969. Pollen record at the Mer Bleue.
Canadian Field-Naturalist 83(1): 7-13.
Corbet, P.S. and G.S. Walley. 1969. The dragonflies
(Odonata) of the Mer Bleue. Canadian Field-Naturalist.
83(1): 14-16.
Dunston, J. 1970. The Mer Bleue bog. Trail & Landscape
4(1): 26-31.
Fassett, N. C. 1957. A manual of aquatic plants. University
of Wisconsin Press, Madison. 405 pp.
Fernald, M. L. 1933. Recent discoveries in the Newfound-
land flora. Rhodora 35: 1-16, 47-63, 80-107, 120-140,
161-185, 203-223, 230-247, 265-283, 298-315, 327-346,
364-386, 395-403.
Freeman, T. N. 1969. Some historical notes on collecting
Lepidoptera in the Mer Bleue. Canadian Field-Naturalist
83(1): 16-18.
Hobson, G. D. 1970. Bedrock features of the Mer Bleue
area by seismic methods. Canadian Field-Naturalist 84(1):
35-38.
Holmgren, P. K.and W. Keuken. 1974. Index herbariorum
Part 1. The herbaria of the world. 6th edition. Regnum
Vegetabile 92: 1-397.
398
Joyal, R. 1971. La tourbiére a sphaignes Mer Bleue prés
d’Ottawa. II. Quelques facteurs écologiques. Canadian
Journal of Botany 50: 1209-1218.
Lafontaine, J.D. 1971. What ever happened to Habenaria
blephariglottis? Trail & Landscape 5(5): 134-136.
Morris, F. 1922. Bladderworts of Ontario. Canadian Field-
Naturalist 36(7): 123-126.
Muenscher, W.C. 1944. Aquatic plants of the United
States. Comstock Publishing Co. Inc., Cornell University,
Ithaca, New York. 374 pp.
Nystrom, E. and S.A. Anrep. 1909. Investigation of the
peat bogs and peat industry of Canada, during the season
1908-1909. Canada Department of Mines, Mines Branch,
Bulletin 1, Report 30.
Ouellet, H., S. J. O'Donnell, and R. A. Foxall. 1976. Gray
Jay nesting in the Mer Bleue Bog, Ottawa, Ontario.
Canadian Field-Naturalist 90(1): 5-10.
Roland, A. E. and E. C. Smith. 1969. The flora of Nova
THE CANADIAN FIELD-NATURALIST
Vol. 93
Scotia. Part 2. The dicotyledons. Proceedings of the Nova
Scotian Institute of Science 26(4): 277-746.
Small, E. 1976. Insect pollinators of the Mer Bleue peat bog
of Ottawa. Canadian Field-Naturalist 90(1): 22-28.
Smith, D. A. 1970a. Notes on bog ecology. Trail & Land-
scape 4(5): 135-143.
Smith, D. A. 1970b. Observations on nesting Hawk Owls
at the Mer Bleue, near Ottawa, Canada. Canadian Field-
Naturalist 84(4): 377-383.
Terasmae, J. and R. J. Mott. 1964. Pollen deposition in
lakes and bogs near Ottawa, Canada. Canadian Journal of
Botany 42: 1355-1363.
Thomson, J. W., Jr. 1940. Preliminary reports on the flora
of Wisconsin. 27. Lentibulareaceae. Wisconsin Academy
of Sciences, Arts and Letters 32: 85-89.
Received 13 February 1979
Accepted 24 April 1979
atl al
Movements of Sympatric Species of Snakes at Amherstburg, Ontario
B. FREEDMAN! and P. M. CATLING2
'Department of Botany, Erindale College, University of Toronto, Mississauga, Ontario L5L 1C6
Present address: Department of Biology and Institute for Resource and Environmental Studies, Dalhousie University,
Halifax, Nova Scotia B3H 4J1
2Department of Botany, University of Toronto, Toronto, Ontario M5S 1A1
Freedman, B. and P. M. Catling. 1979. Movements of sympatric species of snakes at Amherstburg, Ontario. Canadian
Field-Naturalist 93(4): 399-404.
In order to investigate the movements of Butler’s Garter Snakes (Thamnophis butleri), Eastern Garter Snakes ( Thamnophis
sirtalis sirtalis), Brown Snakes ( Storeria dekayi), and Eastern Fox Snakes (Elaphe vulpina gloydi), we systematically searched
for, captured, and individually marked by scale-clipping these species. Recaptured snakes were found to have moved
relatively short distances, with over 50% of marked snakes being recaptured less than 50 m from the initial point of capture
after various recapture intervals. There was no evidence of large-scale emigration of snakes from areas inhabited in the early
spring. For all species, only a small percentage of the population demonstrated relatively long-distance (about 400 m)
unidirectional movements. There were no consistent differences in movements between sexes. More female 7. butleri were
recaptured than males. No directional trends in movements were observed, but a tendency to avoid road-crossing was noted
for T. butleri. In cases of multiple recapture, minimum activity ranges are presented.
Key Words: Thamnophis butleri, Thamnophis s. sirtalis, Storeria dekayi, Elaphe vulpina gloydi, movements, activity range,
southwestern Ontario.
Gregory and Stewart (1975) described dispersal of
the Red-sided Garter Snake (Thamnophis sirtalis
parietalis) following spring emergence from karst
hibernacula in the Interlake District of Manitoba.
There are no other published data on the movements
of snakes in Canada, although these are an important
part of general ecological understanding and are
critical for the conservation of populations of rare
animals.
The purpose of our study was to describe move-
ments of four sympatric species of snakes and to
analyze these with respect to the extent of suitable
habitat required, and the sizes of areas that must be
protected to ensure the survival of populations. The
study area, 2.4km NE of Amherstburg, Essex
County, Ontario, has been described previously
(Freedman and Catling 1978). Two of the species
studied, Butler’s Garter Snake (Thamnophis butleri)
and the Eastern Fox Snake (Elaphe vulpina gloydi)
are considered rare or endangered in Canada (Cook
1970, 1977; Gregory 1977). The other two species, the
Eastern Garter Snake (Thamnophis sirtalis sirtalis)
and the Brown Snake (Storeria dekayi), are wide-
spread and common.
Methods
We searched systematically for snakes along pre-
determined census routes during a series of visits to
the study area in the spring and summer of 1976 (14
and 30 May, 15 June, 10 and 24 July). All snakes
captured were marked by clipping scales (Freedman
and Catling 1978), and the point of capture was
399
recorded on _ large-scale aerial photographs
(1 cm = 10 m). Sex and total length were also record-
ed for each snake to allow comparisons among sexes
and size classes (Freedman and Catling 1978). All
snakes were released where captured within 15 min.
Young snakes less than | yr old and young-of-the-
year (defined by body length) were difficult to mark by
scale-clipping, and were not included. Snakes less
than | yr old were relatively uncommon (Freedman
and Catling 1978).
For all captured snakes the distance between the
recapture and the most recent point of capture was
determined to give a minimum distance travelled. In
cases of more than one recapture of an individual
snake, these minimum distances were also summed to.
give a total minimum distance travelled. In the cases
of multiple recapture, minimum activity ranges were
calculated from the areas enclosed by the joined
points of capture and recapture. We also considered
the possibility that roads could restrict movements of
snakes; we noted whether snakes crossed an open 10-
m-wide gravel road.
Results and Discussion
Out of a total of 250 individuals of T. butleri, 72 T.
s. sirtalis, and 102 S. dekavi, a minimum of 10
recaptures was made per species, and some indi-
viduals were recaptured more than once. All of these
species exhibited a strong tendency for relatively
short-distance movements, with more than 50% of all
marked individuals being recaptured less than 50 m
from the initial point of capture after various intervals
400
of time (Figures 1-3). In general, total distance moved
by all three species increased the longer the time
interval between initial capture and subsequent
recapture(s). This tendency is illustrated for T. butleri
(Figure 1b-d), for which there were 25 recaptures.
For each of the three species, there were no
consistent differences in movement between the sexes
(Table 1), Although female S. dekayi appear to move
less than males, the sample of males is only two. More
female 7. butleri were recaptured than males (Table
()
uJ
a
=)
fS
a
q
(S)
uJ
a
ep)
uj
x
dq
a
2p)
iw
(e)
(e)
2
THE CANADIAN FIELD-NATURALIST
Vol. 93
1),, even though the proportion of males in the
population was about 0.56 (Freedman and Catling
1978). Possibly female 7. butleri are less mobile and
have a greater tendency to bask in the open (Gregory
1975) and are consequently more often captured than
males.
The S. dekayi covering the greatest distances were a
male 31 cm long covering 374 m in 30 d, and a gravid
female 39 cm long covering 226 m in 30 d, both single
recaptures. The 7. butleri covering the greatest
Thamnophis butleri
All recaptures
N=24
LI
Recaptures after |3-I6days
N=7
pay
Recaptures after 30-40 days
Recaptures after 54-70 days
N=1l0
P
in
430-440
510-520
MINIMUM MOVEMENT
FIGURE 1. Minimum movements of marked Thamnophis butleri: (a) all recaptures during 1976; (b,c, d) recaptures at specific
time intervals following release. Data include 20 individuals, of which two were recaptured twice and one was
recaptured three times.
1979 FREEDMAN AND CATLING: DISTANCE MOVEMENTS, SYMPATRIC SNAKES 401
(a)
a5 Thamnophis sirtalis sirtalis
> a, eee
=
ae
dq
Oo
uJ
a
wn > N=12
uJ
x
= 2
(ep)
w
oc |
fe)
2
Oo O O oO O oO O O oO Oo O O O
= aN) no) t wo op) O = (aN) ~m Tt wo oO
Sir ee ee eee ie Ia oe thane ap sh Sli ona
(e) O Oo (e)
FIGURE 2. Minimum movements of marked Thamnophis sirtalis sirtalis from all 1976 recaptures. Data include 11 indi-
viduals, one recaptured twice.
(a)
uJ :
or Storeria dekayi
= par tia Panacea ceo
=
au
6
oO
uJ
rs N=10
op)
uJ
x
<q
2
cop)
w
oO
fe)
2
©) O Oo oO O O O Oo Oo
| I l |
oO oO oO Oo oO oO ! I
= oO O
N DD)
MINIMUM MOVEMENT (m)
FIGURE 3. Minimum movements of marked Storeria dekayi from all 1976 recaptures. Data include eight individuals,
two of which were recaptured twice.
402 THE CANADIAN FIELD-NATURALIST Vol. 93
TABLE |—Sex-specific minimum movements for marked individuals of three species of snakes
Minimum distance travelled (m)
Males Females
Species Mean = SE n Range Mean + SE n Range
Thamnophis butleri 92 +59 5 < 10-433 67 = 28 19 < 10-226
Thamnophis sirtalis Ses 28 6 < 10-147 TD x2 WY 5 < 10-153
Storeria dekayi NG se 7/7) 2 15-374 59 = 34 6 < MOHS7/
distances were also single recaptures, including a 31-
cm male covering 433 m in 70 d and a 47-cm gravid
female covering 517 m in 70 d. The longest distance
moved by 7. s. sirtalis involved a 63-cm female which
covered 153 m in 27 d.
Generally, there were no directional trends in the
movements observed. Thamnophis s. sirtalis moved
only within a restricted and relatively moist area,
where these snakes were abundant. Long-distance
movements of S. dekayi involved moving along a
moist roadside ditch. It might be assumed that the
relative scarcity of T. but/eriin midsummer in the dry
quarry section of the study area was the result of
emigration. The recapture data do not, however,
support this assumption. Of the 17 7. butleri marked
in a dry upland area, 14 were recaptured again in the
immediate vicinity, and eight of these were recaptured
more than 40 d after initial capture. Only three of the
recaptured snakes had moved to lower moist sites.
Two of these, recaptured 70 d after initial capture, had
moved over 400 m to wetter ground. The fact that T.
butleri did not become conspicuously more abundant
in moist places in the western portion of the study area
in midsummer suggests that only asmall percentage of
the population was involved in large-scale emigration.
Carpenter (1952) studied a large population of T.
butleri at a site in Michigan, and found minimum
movements of 26 snakes to average 115 m. He also
found that females moved 65% farther than males. No
directional trends were exhibited in the movements of
snakes in Carpenter’s study.
R. J. Planck and J. T. Planck (1977, unpublished
report, Department of Supplies and Services, Ot-
tawa), in a mark-recapture study of 7. butleri in
southwestern Ontario, found this species to be
relatively sedentary. They found a maximum dis-
placement of only 25 m among nine males, and of
19 m among 19 females. In addition they noted an
apparent lack of directional trend.
Data on movements of two other species of
Thamnophis (T. radix and T. brachystoma), which
have strong taxonomic affinities to 7. but/eri(Conant
1950), may also serve as comparisons with data on the
latter species. Siebert and Hagen (1947), working at a
site in Illinois, found 7. radix to be relatively
sedentary, with 83% of their 41 snake recoveries
showing minimum movements of less than 46 m, and
only one snake moving farther than 90 m. Klingerer
(1957), working with 7. brachystoma in Pennsyl-
vania, also offered evidence for small activity areas,
although Asplund (1963) thought that his later work
provided evidence to the contrary.
Carpenter (1952) recorded movements of T. s.
sirtalis, which were sympatric with 7. butleri at his
Michigan site. He found minimum movements of 127
T. s. sirtalis to average 58 m, about half the distance
averaged by 7. burleri. No differences in movements
were noted between sexes, nor was any directional
trend observed.
Fitch (1965) described movements of T. s. sirtalis in
Kansas, reporting that the minimum distance travel-
led by males averaged 532 m, and by females 347 m.
He also noted a lack of directional trend in the
movements. Fitch also reported that the minimum
distances travelled tended to increase as the time
interval between capture and recapture increased.
Gregory and Stewart (1975), working in the
Interlake region of Manitoba, reported dispersal data
for 26 marked individuals of JT. sirtalis
parietalis. The minimum movements reported
(mean + SE = 10.7 + 0.73 km, range = 4.3 to 17.7 km)
were much longer than those found in the present
study for 7. s. sirtalis. No differences relating to sex
were observed. These authors found the movements
of their marked snakes to be strongly directional
between karst hibernacula and summer feeding
grounds in nearby marshes. The spatial separation of
the hibernacula and the feeding habitat of these
snakes presumably accounts for the relatively long
distances travelled. But no such spatial separation of
hibernacula and feeding areas was apparent at our
site.
Noble and Clausen (1936) reported that little
movement of S. dekayi occurred if captured indi-
viduals were released close to the point of capture;
however, if they were released at another locality, they
tended to return to the original site, moving up to
1200 m in 7 d.
1979
Only one recapture was made by us of a marked
Elaphe vulpina gloydi, out of a total of 33 marked.
This individual, marked in 1976 and recaptured in
1977, travelled a minimum distance of 250 m. Rivard
(1976) reported recapturing 10 E. vulpina gloydi at
Point Pelee National Park, Ontario. These were all
recaptured at or within several metres of their original
point of capture up to 25 d later. Rivard (1976) cited
previous park studies, but noted that they provided
little additional information on E. vulpina gloydi
activity. Fitch (1963) found minimum movements of
up to 1000 m in the congeneric Black Rat Snake
(Elaphe o. obsoleta) at a site in Kansas, although most
recorded movements were less than 200 m.
Various studies have suggested that most snakes do
not wander randomly over the landscape, but rather
inhabit restricted areas, which have been referred to as
“home ranges” or “activity ranges” (Stickel and Cope
1947; Carpenter 1952; Fitch 1963). Stickel and Cope
(1947) calculated that E. 0. obsoleta and the Northern
Black Racer (Coluber c. constrictor) at their sites
could easily cross their home ranges in3 hat anormal
crawling speed. They noted that relatively long-
distance, unidirectional movements do occasionally
occur, but that they represent exceptional events.
Similarly, Hirth et al. (1969) reported that most
recaptures of marked snakes occurred within about
1 km of a hibernaculum in Utah. Noble and Clausen’s
(1936) evidence for homing behavior of S. dekayialso
suggests affinity for a specific area.
The longest distances travelled by snakes in our
study area represent single recaptures, and suggest
relatively long-distance unidirectional movements.
Possibly a small percentage of the population is
comprised of transients. The few cases of two or three
recaptures of a single individual, and minimum
activity range, are summarized in Table 2.
Carpenter (1952) estimated average and maximum
activity ranges of 8000 m2 and 17000 m? for 7. s.
sirtalis, 3000 m2 and 9000 m2 for T. butleri at a site in
Michigan where they were sympatric. Fitch (1965),
working at a site in Kansas, calculated an activity
range of 14.2 ha (142 000 m2) for male 7. s. sirtalis,
and 9.2 ha (92 000 m2) for females. Although Gregory
and Stewart (1975) did not present estimates of
activity ranges for Manitoba Interlake T. sirtalis
parietalis, the long distances travelled by marked
snakes during their dispersal movements from karst
hibernacula (previously described) would suggest very
large activity ranges relative to other observations for
Thamnophis spp. Finally, Fitch (1963) calculated
activity ranges for Elaphe o. obsoleta, a congeneric
with E. vulpina gloydi, and found these to be 11.7 ha
for male snakes and 9.3 ha for females.
Although our data (no crossings of a 10-m-wide
gravel road were observed for five 7. butleri) are too
FREEDMAN AND CATLING: DISTANCE MOVEMENTS, SYMPATRIC SNAKES
403
TABLE 2—Minimum activity ranges of marked snakes
having more than one recapture. Except for the male T. s.
sirtalis, all the snakes were females. Data collected near
Amherstburg, Ontario in 1976
TL, Area of
Species cm Marked Recaptured activity m2
T. butleri 47.0 14 May 30 May 50
10 July
T. butleri Bild 14 May 30 May 50
10 July
T. butleri 47.0 14 May 30 May 600
15 June
24 July
15 8: 50.0 30 May 15 June 2400
sirtalis 10 July
S. dekayi 34.4 30 May 15 June 50
10 July
S. dekayi 39.5 15 June 10 July 1200
24 July
*TL = Total length.
few to be statistically significant, we suspect that there
is a tendency for snakes to avoid crossing roads. The
abilities of both S. dekayi and T. butleri to persist in
isolated parks and vacant lots surrounded by urban
development (Logier 1958; Campbell 1971) suggest
that roads and other habitat discontinuities such as
woodlands (Carpenter 1952) are restrictive to move-
ments of a large proportion of the populations of these
species. This theory deserves more rigorous testing
because it has important implications with respect to
road construction in parks and nature reserves. In
addition to restricting movements, roads may con-
tribute to mortality owing to traffic and to increased
exposure to natural predation.
Acknowledgments
We thank F. R. Cook of the National Museum of |
Natural Sciences, Ottawa, C. A. Campbell of Water-
loo, Ontario, P. T. Gregory of the University of
Victoria, Victoria, and W.F. Weller of Erindale
College, University of Toronto for their critical
reading of the manuscript and their helpful sug-
gestions. The field assistance of S. M. McKay and
K. L. McIntosh is gratefully acknowledged.
Literature Cited
Asplund, K. A. 1963. Ecological factors in the distribution
of Thamnophis brachystoma (Cope). Herpetologica 19(2):
128-132.
Campbell, C. A. 1971. Butler’s Snake in Canada: a review
of previously recorded and newly recorded colonies.
Canadian Amphibian and Reptile Conservation Society
Bulletin 9(5): 1-5.
404
Carpenter, C. C. 1952. Comparative ecology of the Com-
mon Garter Snake (Thamnophis sirtalis), the Ribbon
Snake (Thamnophis sauritus), and Butlers Garter Snake
(Thamnophis butleri) in mixed populations. Ecological
Monographs 22(4): 235-258.
Conant, R. 1950. On the taxonomic status of Thamnophis
butleri Cope. Bulletin of the Chicago Academy of
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Cook, F. R. 1970. Rare or endangered Canadian reptiles
and amphibians. Canadian Field-Naturalist 84(1): 9-16.
Cook, F. R. 1977. Review of the Canadian herpetological
scene. Jn Canada’s threatened species and habitats. Edited
by T. Mosquin and C. Suchal. Canadian Nature Federa-
tion, Special Publication 6, Ottawa. pp. 117-121.
Fitch, H. S. 1963. Natural history of the Black Rat Snake
(Elaphe o. obsoleta) in Kansas. Copeia 1963(4): 649-659.
Fitch, H. S. 1965. An ecological study of the Garter Snake
(Thamnophis sirtalis). Museum of Natural History,
University of Kansas Publications 15: 493-564.
Freedman, W. and P.M. Catling. 1978. Population size
and structure of four sympatric species of snakes at
Amherstburg, Ontario. Canadian Field-Naturalist 92(2):
167-173.
Gregory, P. T. 1975. Aggregations of gravid snakes in
Manitoba. Copeia 1975(1): 185-186.
Gregory, P. T. 1977. Rare and threatened snake species of
Canada. Jn Canada’s threatened species and habitats.
Edited by T. Mosquin and C. Suchal. Canadian Nature
THE CANADIAN FIELD-NATURALIST
Vol. 93
Federation, Special Publication 6, Ottawa. pp. 122-126.
Gregory, P. T. and K. W. Stewart. 1975. Long-distance
dispersal and feeding strategy of the Red-sided Garter
Snake (Thamnophis sirtalis parietalis) in the Interlake
of Manitoba. Canadian Journal of Zoology 53(3): 238-
245.
Hirth, H. F., R.C. Pendleton, A.C. King, and T.R.
Downard. 1969. Dispersal of snakes from a hibernacu-
lum in northwestern Utah. Ecology 50(2): 332-339.
Klingerer, D. 1957. A marking study of Short-headed
Garter Snake in Pennsylvania. Herpetologica 13(2): 100.
Logier, E. B.S. 1958. The snakes of Ontario. University
of Toronto Press, Toronto. 94 pp.
Noble, G. K. and H. J. Clausen. 1936. The aggregation
behavior of Storeria dekayi and other snakes, with
especial reference to the sense organs involved. Eco-
logical Monographs 6(2): 269-316.
Rivard, D.H. 1976. The biology and conservation of
Eastern Fox Snakes (Elaphe vulpina gloydi). M.Sc.
thesis, Carleton University, Ottawa. 64 pp.
Siebert, H. C. and C. W. Hagen. 1947. Studies on a popu-
lation of snakes in Illinois. Copeia 1947(1): 2-22.
Stickel, W.H. and J.B. Cope. 1947. The home ranges
and wanderings of snakes. Copeia 1947(2): 127-136.
Received 13 February 1979
Accepted 8 June 1979
Winter Feeding by Porcupines in Montane Forests of
Southwestern Alberta
LAWRENCE D. HARDER!
Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9
!Present address: Department of Zoology, University of Toronto, Toronto, Ontario M5S 1A1
Harder, L. D. 1979. Winter feeding by Porcupines in montane forests of southwestern Alberta. Canadian Field-Naturalist
93(4): 405-410.
Past feeding on the inner bark of Douglas Fir (Pseudotsuga menziesii) and Limber Pine (Pinus flexilis) by Porcupines
(Erethizon dorsatum) was examined on two areas of southwestern Alberta. Measurements of annual growth from increment
cores were used to estimate the previous sizes of trees. In conjunction with the ages of feeding scars, these estimates provided a
means of determining the physical characteristics of the trees used by Porcupines over a 20-yr period. Of the three community
types available to Porcupines, pure stands of Douglas Fir in leeward situations were preferred. Although Porcupines fed in
the larger, faster growing trees within this community on both study areas, the average diameter of the boles of used trees
differed between study areas. This may be related to a possible relaxation of the preference for larger trees on the more
extensively used area.
Key Words: Porcupine, Erethizon dorsatum, food habits, Alberta.
Concurrent with the cessation of plant growth and
the onset of breeding in autumn, Porcupines ( Erethi-
zon dorsatum) move to areas more heavily wooded
than their summer ranges and begin to feed on the
inner bark of woody plants and the foliage of several
species of conifers (Gabrielson and Horn 1930; Curtis
and Wilson 1953; Brander 1973). When feeding on
inner bark, Porcupines eat the cambium and phloem
and expose the underlying xylem. During subsequent
growth periods, exposed xylem is slowly covered by
the centripetal growth of callus tissue formed from the
surrounding cambium (Brown 1971). The resulting
scars are a semi-permanent record of the amount and
location of bark removed and the annuli in the new
xylem relate the year in which feeding occurred.
Only two published studies (Spencer 1964; Storm
and Halvorson 1967) have taken advantage of the
temporal information available in feeding scars;
neither were directly concerned with the food habits of
Porcupines. I have studied the distribution of feeding
scars within and among trees to determine the general
pattern of forest use by Porcupines on two study areas
in southwestern Alberta. This report documents the
physical characteristics of the trees used by Porcu-
pines during a 20-yr period, and the location of
feeding within those trees.
Study Areas and Methods
Two study areas were examined, both located on
the eastern fringes of the foothills of the Front Ranges
of the Rocky Mountains. The Zoratti study area
(49°27’N, 114°04’W), 11: km W of Pincher Creek,
Alberta, covers an area of 36.3 ha and varies in
elevation from 1210 to 1320 m. The Skelton study
area (49°34’N, 114°13’W), 16 km NW of the Zoratti
site, encompasses 115.4 ha and ranges in elevation
from 1180 to 1360 m.
The frequency and intensity of prevailing westerly
winds in this region influence the distribution and
composition of resident plant communities (Moss
1944). Ridgetops and windswept western exposures
are occupied by an open, Limber Pine ( Pinus flexilis)—
Douglas Fir (Pseudotsuga menziesii var. glauca)
woodland (windward community). More protected
north- and east-facing slopes are dominated by stands
of Douglas Fir (leeward pure community) with
occasional hybrid spruce (Picea glauca X P. engel-
mannii, see Daubenmire 1974). The gentler leeward
slopes at the base of the ridge that comprised the’
Zoratti area are inhabited by mixed stands of Limber
Pine and Douglas Fir (leeward mixed community)
which appear to be seral to the adjacent leeward pure
community.
The coniferous communities were sampled in
20 X 20 m plots, but the sampling intensity differed on
the two study areas. Approximately 10% of the
Zoratti site was examined with 38 plots whose centers
were 45 m apart along transects at 100-m intervals.
The Skelton area was too large to be sampled this
intensively and 40 plots were chosen from an original
190, distributed as on the Zoratti area, on the basis of
community type, tree density, and the uniformity of
tree distribution. Several plots that did not include
more than 20 trees were enlarged to cover an area of
500 m’.
405
406
Species, circumference at breast height, height, and
evidence and the probable cause of any leader damage
were determined for all trees taller than breast height
(1.35 m) within each sample plot. A random selection
of 20% of these trees was aged with an increment
borer. Each tree was inspected for feeding scars and a
record made of the intensity of use (four classes based
on total scar area) for the tree as a whole and
separately for lateral branches and the bole. The
number of years since feeding occurred, the height
range, and the intensity of use for each year of feeding
were estimated. I determined year of feeding by
removing a portion of the scar tissue surrounding the
feeding area and counting the annual rings from the
level of exposure by the Porcupine to the current
xylem surface.
Limber Pine and Douglas Fir are considered
excellent dendrochronological material because of a
high correlation between ring formation and the
annual climatic cycle (Schulman 1956). The age of
very old scars on senescent branches at the base of the
crown, however, was probably measured less ac-
curately than the age of the tree because these old
branches produce xylem rings irregularly (Reukema
IPS).
Several characteristics were calculated for each
tree. Diameter at breast height (DBH) was derived
from the circumference. The ratio of DBH to height
served as an index of growth form based on the direct
relation between bole diameter and crown width
(Mitchell 1969). The magnitude of this confirmation
index (CI) generally ranged from 0.75 to 3.00 and was
useful only for intraspecific comparisons because
growth form varies between species. Estimates of the
average annual growth of the bole and height of the
tree were calculated by dividing DBH and height,
respectively, by age.
Because current DBH is only indirectly related to
the dimensions of trees considered by Porcupines
during previous years, I estimated the DBH and
height of each tree for the preceding 20 yr. Annual
growth increments (+5 X 10° mm) for each of the
past 20 yr were measured for each core with a
dissecting microscope. The length of each increment
core, from the pith to its outer surface, and the
thickness of the bark (+0.025 mm) extracted at breast
height with the increment borer, were measured with
vernier calipers.
Calculation of DBH, x yr prior to measurement,
employed the following basic algorithm. The radius
inside the bark during the year of interest (IRy) was
taken as the product of the inside radius at the time of
measurement ((DBH/2)-BARK) and the proportion
of the total length (TC) represented by the length of
the core x yr previous (TC- 2 1, ;where I, is the length
of the growth increment for any given year, n. There-
THE CANADIAN FIELD-NATURALIST
Vol. 93
fone; i
IR, = ((DBH/2)-BARK) ° (TC- 21 )/TO),
and the DBH during that year was estimated with the
linear regression equation that related the inside
radius at the time of measurement to the measured
DBH for a particular species in a given community
(Husch et al. 1972). The error associated with this
regression equation is probably negligible since the
smallest coefficient of determination (r’) was 0.997.
Regression equations that described the relation
between the calculated DBH and the age and
measured DBH were computed for each of the 20 yr
preceding measurement from the data for cored trees
from each forest type and were used to estimate past
diameters for uncored trees.
The procedure applied in estimating the height of a
tree during each of the 20 yr prior to measurement,
took advantage of the relation between height and
DBH. The growth ofa tree was taken asa straight line,
on a log-log plot, between the measured height and
DBH and a height of 1.35 m and zero DBH, which
allowed the estimation of the height corresponding to
any previously established DBH. If primary growth
had been interrupted by a factor other than that of a
Porcupine killing the leader, or if the tree was girdled
more than 20 yr before measurement, previous
heights were not calculated. If the leader was killed by
Porcupines within 20 yr of measurement, the height
during any year since the feeding occurred was
estimated from a regression line between the
measured height and DBH, and the height of the
uppermost feeding scar and the calculated DBH for
the year during which feeding occurred.
With the exception of leeward pure forests, only a
small proportion of the available trees was used by
Porcupines during a given year and the average
position of each tree in relation to the community
mean during the 20 yr prior to measurement, has been
examined rather than the tree’s absolute size or age.
Data for DBH, height, and CI were normalized
([Xi—X])/s; Sokal and Rohlf 1969) for each of the
20 yr prior to measurement. The average trans-
formation of each of the three variables for each tree
during this period, therefore, becomes an indicator of
its relative status in the community, measured in
standard deviations from the mean of zero.
Results
Preferences for Individual Trees
Porcupines fed on 37.1% (368) of the 993 Douglas
Fir and Limber Pine sampled on the Zoratti area.
Only 17.8% (78/439) of the conifers in the windward
community were used by Porcupines, while 24.4%
(40/164) of the trees in the leeward mixed community
and 64.1% (250/390) of the trees in the leeward pure
community were used. A significantly smaller pro-
1979 HARDER: TREES UTILIZED BY PORCUPINES
407
TABLE |—Comparisons of the ages, relative sizes, and estimated annual growth increments of trees fed upon by Porcupines with unused trees on
the Zoratti and Skelton areas. Limber Pine in the Skelton windward community were not compared because only two trees were
used during the 20 yr preceding measurement. Means +SE (n) are given. Species/ location code: first character Z—Zoratti or S—Skelton:;
second character W—windward, P—leeward pure, or M—leeward mixed; third character F—Douglas Fir or P—Limber Pine
Age DBH growth Height growth
Species (yr) DBH Height Cl (cm/ yr) (m/ yr)
location Used Unused Used Unused Used Unused Used Unused Used Unused Used Unused
ZWF 60.2 59.1 0.57* -0.28 0.30 -0.21 0.58 0.34 0.18 0.14 0.08 0.07
+11.55 +4.96 +0.403 +0.119 +0.368 +0.129 +0.475 +0.120 +0.048 +0.019 +0.018 +0.005
(9) (17) (10) (57) (10) (56) (10) (56) (9) (17) (8) (17)
ZPF 60.8 58.3 0.19*** 0.37 0.14*** -0.30 0252" * 0.45 0.29** 0.22 0.17 0.15
+2.09 +3.51 +0.070 +0.074 +0.069 +0.088 +0.074 +0.076 +0.015 +0.018 +0.008 +0.009
(70) (44) (205) (176) (182) (159) (182) (159) (70) (44) (52) (43)
ZMF 62.6*** 33.9 1.08*** -0.42 1.00*** ~0.38 0.70** ~0.43 0.34** 0.19 0.15 0.13
263} 5 +4.13 +0.315 +0.107 +0.311 +0.113 +0.239 +0.131 +0.030 +0.031 +0.016 +0.012
(12) (16) (9) (59) (9) (58) (9) (58) (12) (16) (10) (15)
ZWP 73.2 66.5 0.32 0.08 0.31 -0.08 0.18 0.08 0.15* 0.12 0.05 0.04
+8.08 Staley 2. +0.217 +0.050 +0.183 +0.058 +0.179 +0.060 +0.013 +0.006 +0.004 +0.002
G1) (81) (47) (325) (45) (266) (45) (266) (1) (81) (21) (74)
ZMP 62.4* 44.1 0.88*** -0.27 0.62* -0.25 O93 es* -0.33 0.29 0.21 0.10 0.11
+7.30 +3.65 +0.254 +0.107 +0.210 +0.125 +0.316 +0.177 +0.035 +0.023 +0.013 +0.007
(7) (23) (11) (85) (10) (69) (10) (69) (7) (23) (5) (20)
SWF 91.0** 70.8 0.91*** -0.07 0.77** -0.07 0.35 -0.03 0.30 0.21 0.12 0.09
+6.77, +2.41 +0.253 +0.063 +0.214 +0.066 +0.193 +0.068 +0.056 +0.014 +0.022 +0.006
(9) (45) (18) (236) (18) (220) (18) (220) (9) (45) (7) (42)
SPF DS ee, 63.8 0.83*** -0.18 0.34*** -0.11 0.97*** -0.21 0:33%%* 0.26 0.20** 0.17
+1.88 ae ||72}| +0.079 +0.029 +0.089 +0.033 +0.091 +0.029 +0.019 +0.007 +0.011 +0.004
(45) (189) (163) (1058) (151) (977) (151) (977) (45) (189) (25) (184)
*P< 0.05.
**P< 0.01.
*** P< (2.001.
portion (22.4%: log likelihood ratio, P< 0.005) of the
1601 conifers sampled on the Skelton area were fed
upon by Porcupines; windward trees there were also
used less frequently (12.3%, 45/367) than trees in the
adjacent leeward pure community (25.4%, 313/ 1234).
During the 20 yr preceding measurement, Porcu-
pines usually fed in trees that were significantly larger
and had more extensive crowns than the trees they
neglected, even though the selected trees were not
necessarily statistically older than unused trees (Table
1). Inaddition, Douglas Fir fed upon by Porcupines in
the preferred leeward pure communities grew
significantly faster, at least in bole diameter, than
unused trees (Table 1). Porcupines on the Zorattiarea
also used Douglas Fir in the leeward mixed
community, and Limber Pine in the windward
community that grew faster in bole diameter than the
trees they neglected. Stepwise discriminant analyses
(Table 2) indicate that DBH and growth form are
more effective characteristics for distinguishing used
TABLE 2—Discriminant analysis statistics detailing variables that best distinguished between trees fed upon by Porcupines
and unused trees on the Zoratti and Skelton areas!
Species, Independent Discriminant Wilks’
location? variable} Constant coefficient lambda
ZWF CI (56,10) 0.194 0.952 0.900**
ZPF CI (159,182) 0.077 0.968 0.887***
ZMF DBH (58,9) 0.212 1.023 0.728***
ZWP DBH (266,45) 0.026 0.959 0.976**
ZMP CI (69,10) 0.164 0.944 0.841***
SWF DBH (220,18) —0.004 1.018 0.931***
SBE CI (997,151) 0.051 0.971 0.847***
'Discriminant analyses were first performed on aged trees only, but because age was not an effective discriminator the analyses presented are
based on all trees.
2Species/location code given in Table 1.
3Quantities in parentheses represent the number of unused and used trees during the 20-yr period preceding measurement.
**P< 0.01.
*** P< 0.001.
408 THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 3—Mean =+SE (n) estimated size of Douglas Fir in leeward pure communities during 1955 and 1970, and of trees
fed upon by Porcupines, 1954-1973
1955
Zoratti area
DBH (cm) 11.6 = 0.38 (343)
Height (m) 7.6 = 0.18 (313)
Cl 1.41 + 0.033 (309)
Skelton area
DBH (cm) 17.1 = 0.26 (1134)
10.8 + 0.11 (1050)
Height (m)
Cl 1.55 + 0.017 (1041)
trees from unused trees than age or height. Any
tendency for used trees to be older or taller was
probably an artifact of the strong intercorrelations
between all four variables.
Only in the leeward pure forests were trees fed upon
with sufficient frequency to allow an examination of
the absolute size of used trees. Trees used during the
20 yr prior to measurement in the Skelton leeward
forest were significantly larger than those used on the
Zoratti area (t-test, DBH P< 0.001; height
P< 0.001; CI P< 0.001). Although trees on the
Skelton area were generally larger than those on the
Zoratti area (Table 3), this difference is not sufficient
to explain the disparity in the size of trees used.
Location and Amount of Feeding within Trees
Although Porcupines feed on both the inner bark
and foliage of conifers, evidence of feeding on foliage
is ephemeral and cannot be assessed after several
years. Porcupines tend to feed in the upper portions of
conifers on the bole and lateral branches (Tables 4, 5;
Gabrielson 1928: Taylor 1935; Shapiro 1949). Feeding
1970 Used
16.3 = 0.50 (381)
9.2 + 0.22 (341)
1.70 + 0.034 (341)
16.9 = 0.55 (256)
Dei ae Py (73)
1.81 + 0.045 (232)
19.4 + 0.30 (1211)
(6 22 0) JIS) (U227A)
1.62 = 0.017 (1117)
26.0 = 0.73 (195)
12.0 + 9.35 (179)
2.22 + 0.054 (179)
scars on the upper surfaces of branches were usually
encountered over a greater height range within a
particular tree than were scars on the bole.
Usually less than 650 cm? of bark were removed
from an individual tree per winter, although many
trees were used more extensively (Table 6). Feeding
most commonly occurred on portions of the tree
unprotected by corky bark, such as the distal ends of
branches and the upper bole. Porcupines often
succeeded in removing bark from the entire cir-
cumference of the bole. This caused the death of the
upper crown and often altered the tree’s form,
particularly if the tree was girdled again in subsequent
years (Table 7).
Discussion
A preference by Porcupines for relatively large fast-
growing trees has been widely reported (Rudolf 1949;
Curtis and Wilson 1953; Krefting et al. 1962; Spencer
1964) and was also evident on the Zorattiand Skelton
areas in this study. Vigorous trees with large open
crowns produce a larger annual increment of phloem
TABLE 4— Mean + SD (n) position of Porcupine feeding within trees on the Zoratti and Skelton areas. Except for the column
on the extreme right, position is measured ona scale on which ground level has a value of 0 and the top of the tree has a value of
1. Position of the mode within the feeding range involved a similar scale on which the bottom and top of the feeding range
within a tree had values of 0 and 1, respectively
Top Bottom
Species, of feeding of feeding
location! range range
ZWFE 0.51 + 0.204 (11) 0.29 + 0.212 (11)
74 NE 0.80 + 0.200 (233) 0.37 + 0.203 (233)
ZMF 0.58 + 0.275 (244) 0.24 + 0.170 (24)
ZWP 0.63 + 0.275 (49) 0.33 + 0.242 (49)
ZMP 0.66 + 0.249 (16) 0.42 + 0.199 (16)
SWF 0.78 + 0.176 (19) 0.37 + 0.199 (19)
Se 0.82 + 0.197 (179) 0.49 = 0.222 (179)
SWP — (2) — (2)
'Species/location code given in Table 1.
Mode
of feeding
within tree
0.38 + 0.207 (9)
0.65 + 0.190 (121)
0.40 = 0.214 (16)
0.51 + 0.237 (40)
0.57 = 0.109 (7)
0.67 + 0.171 (10)
0.66 + 0.180 (78)
= (2)
Mode
of feeding
within
feeding range
0.62 + 0.243 (9)
0.59 + 0.181 (165)
0.60 + 0.164 (13)
0.57 + 0.242 (40)
0.53 + 0.182 (12)
0.72 + 0.171 (19)
0.59 + 0.244 (182)
0.67 + 0.313 (6)
1979
TABLE 5—Relative frequency of feeding on boles and
branches by Porcupines. These data are from trees that were
used during a single year only
Branch Bole
Species, use use
location! greater Equal greater
ZWF 5 0 5
ZPF 96 5 116
ZMF 12 2 7
ZWP 19 | 38
ZMP 3 | 13
SWF . 25 3 8
SPF 208 5 77
SWP | 0 3}
'Species/location code given in Table 1.
TABLE 6—Number of trees and area of bark removed by
Porcupines per year of feeding
S . Area (cm2)
pecies,
location’ < 650 650-1900 1900-3200 > 3200
ZWE 12 0 0 0
ZPF 157 132 8 2
ZMF 16 1] 0 0
ZWP 52 16 3 0
ZMP lS 7 l 0
SWE 38 4 0 0
SPF 300 85 9 2
SWP 6 2 0 0
'Species/location code given in Table 1.
TABLE 7—Frequency of girdling of conifers by Porcupines
on the Skelton area
Frequency of leader death
Species,
location! 0 ; | 2 3
SWF 29 8 0 0
SPF 172 105 30 4
SWP 6 3 0 0
'Species/location code given in Table |.
(Bannan 1955; Grillos and Smith 1959), and provide
more foliage and a greater volume of inner bark than
crowded trees. Because Porcupines generally feed on
inner bark that is accessible from positions that can be
maintained without undue exertion (Taylor 1935;
Spencer 1964), open-crowned trees, which provide
more large branches, allow Porcupines greater access
to food. The phloem and foliage of conifers contain
the greatest concentrations of fats and carbohydrates
HARDER: TREES UTILIZED BY PORCUPINES
409
of the aerial tissues (see Kramer and Kozlowski(1960)
for a review), and the tendency of Porcupines to feed
on these structures within the tree crown where
growth is most rapid may result in the consumption of
the most nutritious food available.
The quality of a food item is a complex property
that involves not only its energy and nutrient content,
but also the digestion and assimilation efficiency of its
consumers (Longhurst et al. 1968), and the influence
of secondary plant compounds (Freeland and Janzen
1974). In addition, an animal’s response to a
particular stimulus, such as food with a specific
nutrient content, can be greatly modified by its
physiological state (Cabanac 1971). An awareness by
Porcupines of chemical qualities is not required to
explain the observed preferences for particular trees,
because vigorous trees may be identified by obvious
physical features. Chemical attributes, however, may
be important within trees because Porcupines appear
to select food items through olfaction (Murie 1926;
Taylor 1935).
The average diameters of used trees from both
study areas (Table 3) fall within the range of size
preferences reported for Porcupines from various
regions of North America (Taylor 1935; Curtis and
Wilson 1953; Krefting et al. 1962; van Deusen and
Myers 1962). A predilection for a particular size class
irrespective of geographic location or tree species
lends support to the suggestion of Curtis and Wilson
(1953) that Porcupines may be most adept at climbing
trees with a DBH between 15 and 25 cm. Presumably
trees can become too large to be readily climbed
(Taylor 1935; Curtis and Kozicky 1944; Curtis and
Wilson 1953), resulting in a decline in their use by
Porcupines even though they may contain large
quantities of high-quality food. Local differences in
the size of used trees of the magnitude observed
between the Zoratti and Skelton areas would not be
expected, however, if Porcupines exercised size
preferences solely related to the ease of climbing trees,
unless the preferred size was unavailable. Storm and
Halvorson (1967) have presented convincing evidence
that the removal of bark by Porcupines greatly affects
the vigor of a tree for up to 10 yr. If a forest, such as
the leeward pure community on the Zoratti area, was
heavily used for several years, the number of rapidly
growing trees within the preferred size class would be
reduced. Porcupines feeding in this forest during
subsequent winters could, therefore, face a choice
between slower-growing trees of the appropriate size
or smaller vigorous trees.
Consumers cannot afford to be selective when
resources are limited (Emlen 1966, 1968). Relaxation
of selectivity may have occurred on the more
extensively used Zoratti area and resulted in the
apparent preference for smaller trees. The trees used
410
on the Skelton area consequently may provide a better
indication of the physical qualities of trees preferred
by Porcupines.
Acknowledgments
This study was supported in part by the National
Research Council of Canada. J. O. Murie and F. C.
Zwickel kindly reviewed various drafts of this paper.
Barney and Anne Zoratti, and Bill and Mary Skelton
generously consented to my activities on their lands.
V. A. Loewen and V. Aziz assisted with the collection
of data on the Skelton area.
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Vol. 93
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Received 3 February 1979
Accepted 29 May 1979
Interactions between Snowy and Short-eared Owls in Winter
M. Ross LEIN and PETER C. BOXALL
Department of Biology, University of Calgary, Calgary, Alberta
T2N IN4
Lein, M. Ross and Peter C. Boxall. 1979. Interactions between Snowy and Short-eared Owls in winter. Canadian Field-
Naturalist 93(4): 411-414.
Observations on habitat use, food habits, and behavioral interactions of Snowy Owls (Nyctea scandiaca) and Short-eared
Owls ( Asio flammeus) in winter near Calgary, Alberta, suggest that no serious competition occurs between them. They differ
in habitat use, obtaining different types of prey as a result. Kleptoparasitism by Snowy Owls upon Short-eared Owls may be
the most frequent interspecific interaction. Reports of Short-eared Owls in the diet of Snowy Owls probably reflect
Opportunistic feeding, rather than a mechanism to reduce competition for food.
Key Words: Snowy Owl, Nyctea scandiaca, Short-eared Owl, Asio flammeus, habitat use, food habits, kleptoparasitism,
Alberta, winter.
Levin et al. (1977) described a case of probable
predation by the Snowy Owl ( Nyctea scandiaca) upon
the smaller Short-eared Owl (Asio flammeus), and
considered the complex relationship between two
possible competitors that may also interact as
predator and prey. They suggest that, besides ob-
taining food, the larger raptor may benefit by
eliminating a potential competitor. We report find-
ings that clarify the interactions between these species
in winter.
Our observations were made near Calgary, Alberta
(SI°N, 114° W), where Snowy Owls regularly winter
(Bird 1972). Short-eared Owls breed near Calgary,
and some overwinter in certain years (Salt and Salt
1976). Hence the two species sometimes come into
contact, providing opportunities to witness inter-
actions.
Methods
Our study area 1s a 185-km? block of agricultural
land southeast of Calgary. Approximately 70% of the
area is cultivated for cereal grains, and about 20% is
hayfields and pasture.. Numerous small sloughs
contain water in spring but support a dense cover of
grasses, sedges, and weeds during the rest of the year.
During visits to the area we recorded the habitat in
which each owl was located when first sighted. If an
owl was on a boundary between two habitats a value
of 0.5 owl was assigned to each.
We analyzed pellets obtained at known roosting
sites and around hunting perches. Most pellets were
readily assigned to one or the other species by
differences in size and by association with known
perches. The identity of a few pellets was question-
able; these were omitted from the analysis. Frag-
mented remnants of an estimated four to six pellets
from a Short-eared Owl roost were also analyzed.
Observations of interspecific interactions were
made whenever possible from 1973 through 1978.
Between November 1977 and March 1978 over 130 h
of behavioral observations of Snowy Owls were made.
Results
The habitat distributions of sightings of the two
species (Table |) are not totally comparable because
of behavioral differences. Many of the Snowy Owl
TABLE |—Habitat distributions of Snowy and Short-eared Owl sightings on the study area in the winter of 1977-1978, plus the
habitat distribution of prey-capture attempts by Snowy Owls
Number of cases (% total)
Habitat Snowy Owl Short-eared Owl Snowy Owl
sightings sightings hunting attempts
Stubblefield 87.5 (43.8) 30.5 (51.7) 14.5 (51.7)
Summerfallow 60.0 (30.0) I (5) 0.5 (1.8)
Hayfield 28.5 (14.3) 7.0 (11.9) 7.0 (25.0)
Pasture 14.0 (7.0) IS (25) 1.0 (3.6)
Ungrazed grassland 6.0 (3.0) ND (loz) 1.5 (5.4)
Slough 4.0 (2.0) 17.5 (29.7) 3.5 (12.5)
Total 200.0 (100.1) 59.0 (100.0) 28 (100.0)
4}
412
sightings probably were of non-hunting birds, be-
cause they roost in conspicuous locations such as on
poles or hilltops. Short-eared Owls use communal
roosts in winter, in locations offering protection from
both weather and easy detection (Clark 1975; personal
observation). Therefore, most Short-eared Owls seen
were flying and probably were hunting. The distri-
bution of prey-capture attempts by Snowy Owls
(Table 1) is also not fully comparable with the Short-
eared Owl sightings because we cannot assume that
all the Short-ears were hunting.
Both species were observed most often in stubble-
fields, which comprised 37% of the study area. While
most hunts by Snowy Owls were also in this habitat
(Table 1), many of the Short-eared Owls recorded in
stubble actually appeared to be hunting over road
ditches or sloughs and were assigned to stubble only
because they were crossing that habitat when first
sighted. Data on prey-capture attempts by Short-
eared Owls are unavailable, but would probably
indicate a lesser importance of this habitat than it has
for Snowy Owls.
The high number of Snowy Owls sighted in
summerfallow relates to their roosting habits, des-
cribed above. They rarely hunted in this habitat
(Table 1). Sloughs, comprising only 3.6% of the study
area, are favored hunting locations for Short-eared
Owls, which frequently are seen coursing back and
forth over them. In contrast, few hunting attempts by
Snowy Owls were observed around sloughs (Table 1).
Table 2 lists the prey items recovered from pellets.
The proportions of Deer Mice (Peromyscus manicu-
latus) and Meadow Voles ( Microtus pennsylvanicus) in
the diets of the two species differ significantly (chi-
square test, P<0.001). Microtus comprised 79% of
prey individuals in Short-eared Owl pellets but were
only half as frequent as Peromyscus in Snowy Owl
pellets.
Remains (feet and skull fragments) of Short-eared
Owls were recovered from two Snowy Owl pellets
found on Snowy Owl territories adjacent to Short-
eared Owl roosts. The range of prey size is greater for
the Snowy Owl, as expected from its larger body size
(Schoener 1969).
Although both Snowy and Short-eared Owls were
seen repeatedly in the same areas, only a few actual
encounters were witnessed. These are summarized as
follows:
1) 18 February 1975. At 15:57 (MST) a Snowy Owl (SO)
flew from a power pole toward a Short-eared Owl (SEO)
hunting 300-400 m away. As it approached, the SEO dove at
it, coming within 2-3 m. The SO landed and assumed a
crouched defensive posture with the body axis at 15-20°
from the horizontal, plumage ruffled, and wings partly
spread. The SEO circled, making repeated shallow stoops,
for 2-3 min before flying away.
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 2—Prey of Snowy Owls and Short-eared Owls on the
study area in the winter of 1977-1978, as determined from
examination of pellets
Number of individuals
(% total)
Prey species Snowy Owl! Short-eared
Owl?
Mammals
Peromyscus maniculatus 79 (51.0) 8 (16.7)
Microtus pennsylvanicus 39 (25.3) 38 (79.2)
Spermophilus richardson? WZ (HT) 0
Mustela frenata* 3) (9) 0
Birds
Perdix perdix> 14 (9.0) 0
Asio flammeus A (V3), —
Passerine spp. 6 (3.9) Di (AD)
Total 155 (100.1) 48 (100.1)
‘Based on analysis of 72 pellets.
2Based on analysis of 22 pellets plus numerous fragments.
3Richardson’s Ground Squirrel.
4Long-tailed Weasel.
5Gray Partridge.
2) 1 December 1977. A SO was perched on a powerline
tower ina pasture. At 16:07 a SEO, flying high above, dove at
it, coming within 2 m. It circled about 10 m above the SO,
then stooped again. There was no observable response. The
SEO then flew off and began hunting ina field 800 m away.
3) 6 December 1977. At 13:45, a hunting SEO landed ina
stubblefield 200 m from a perched SO. The SO flew and
supplanted the SEO, which then circled 8-10 m above it,
uttering “keeow” calls for 20-30 s before flying away. The
SO was not disturbed by this, but appeared to be watching
another SO flying in the distance.
4) 14 December 1977. A SO was perched ona knoll ina
stubblefield. At 13:32 a SEO flew towards it and stooped
several times within | m of the perched SO, which ducked
while watching its attacker. The SEO circled several times,
then flew to a fencepost 900 m away. It resumed hunting at
13:41.
5) 21 December 1977. At 15:00 a SO flew to a power pole
near a SEO that was hunting in a stubblefield. As the SO
landed, the SEO flew toward it and dove twice, coming
within 3 m. Three other SEOs immediately joined the first
and circled 25-30 m above the SO. After | min the SEOs
scattered and resumed hunting. No reactions by the SO were
seen.
6) 21 December 1977. A SO was sitting on the ground
100 m from a slough where a SEO was hunting. Between
17:25 and 17:30 the SEO made three unsuccessful pounces at
prey, without landing for longer than 10 s. The SO watched
the SEO but did not appear aroused. At 17:30 the SEO
landed in the grass, whereupon the SO flew and supplanted
it. The SEO took flight, circled briefly 10-15 m above the
SO, and flew away. The SO then flew back and forth across
179
the slough several times and landed on a fencepost about
150 m away.
7) 19 December 1978. At 15:00 a SO flew from a power
pole toward a SEO flying, with a mouse in its feet, about
500 m away. As the SO approached, both owls disappeared
behind a low rise. Almost immediately the SEO rose straight
up to about 15m with empty feet. It circled the area,
stooping toward the ground several times, before flying
away. The SO, which presumably was perched on the ground
with the mouse, was not re-sighted.
Discussion
The interspecific differences in habitat use and diet
are closely interrelated. Snap-trapping surveys at the
beginning of the winter of 1977-1978 indicated that
Microtus were most common in sloughs and roadside
ditches (Boxall, unpublished data), where Short-eared
Owls frequently were observed hunting. The pref-
erence of Microtus for dense cover is well-known
(e.g., Birney et al. 1976). Peromyscus were more
abundant than Microtus in stubblefields and hay-
fields, where most Snowy Owl hunting attempts were
observed (Table 1).
The failure of Snowy Owls to prey more heavily on
Microtus may be related to hunting behavior. Snowy
Owls are “sit and wait” predators, launching hunting
attempts from perches after prey has been detected
(Hohn 1973; personal observation). In contrast,
Short-eared Owls hunt primarily by flying back and
forth low over vegetation (Clark 1975), pouncing on
any prey located. A small mammal moving within the
dense cover in sloughs or ditches would be screened
from a Snowy Owl perched some distance away, but
would be visible to a flying Short-eared Owl looking
directly down into the vegetation.
The data on habitat use and food habits suggest that
considerable ecological segregation exists between the
two species and that interspecific competition for food
is unlikely under the conditions of our study. Our
behavioral observations support this conclusion.
Snowy Owls initiated four interactions. Two (see
examples 3 and 6) were in apparent response to a
Short-eared Owl landing after prey-capture attempts
that might have been successful. In observation 6 the
Snowy Owl did not react to the Short-ear hunting
nearby until, after 5 min, the latter landed for longer
than a few seconds. We suggest that these actions by
Snowy Owls were attempted piracy, triggered by an
apparently successful hunt by a Short-eared Owl.
Observation 7 appears to be a definite example of
kleptoparasitism. Duffy et al. (1976) describe klepto-
parasitism by a Snowy Owl ona Marsh Hawk (Circus
cyaneus) and Tullock (1968) reported Snowy Owls
robbing Hooded Crows (Corvus corone).
Snowy Owls supplanted Short-eared Owls in
several cases but did not pursue them. These probably
LEIN AND BOXALL: OWL INTERACTIONS
413
were not attempts at predation because Snowy Owls
will pursue flying prey (Meinertzhagen 1959: Nero
1964). Similarly, interspecific territoriality is an
unlikely explanation for such encounters because
Snowy Owls ignored Short-eared Owls unless the
latter appeared to have captured prey. Snowy Owls
are territorial in winter (Keith 1964), but the postures
and modified flight patterns seen in intraspecific
territorial encounters (personal observation) were
absent in these interspecific interactions.
The encounters initiated by Short-eared Owls (see
examples 2, 4, 5), and the reaction of the Short-eared
Owls in observations | and 7, are examples of the
mobbing response of many birds to predators (Hinde
1970). Once mobbing by one Short-eared Owl
attracted three conspecifics to join in briefly. The
“keeow” vocalization noted in observation 3 is an
alarm call of the Short-eared Owl (Clark 1975). The
displays used by Short-eared Owls in intraspecific
territorial encounters (Lockie 1955; Clark 1975) were
not seen. Breeding Short-eared Owls direct such
displays towards various raptors (Clark 1975), and we
have seen a wintering Short-ear display to a Rough-
legged Hawk (Buteo lagopus). Thus the behavior of
Short-eared Owls towards Snowy Owls is an alarm
response rather than an aggressive or territorial one.
The posture of the Snowy Owl while being mobbed in
observation | resembles that of owls attacked by
jaegers (Stercorarius spp.) (Tullock 1968) or by a
Goshawk (Accipiter gentilis) (personal observation).
The lack of territoriality between Snowy Owls and
Short-eared Owls in winter, along with differences in
habitat use and diet, suggests little competition
between them. Short-eared Owls winter in areas with
high populations of small mammals (Clark 1975),
suggesting that winter contact with Snowy Owls will
occur primarily in situations where high food density
alleviates competition (see also Lack 1946). Pitelka et
al. (1955) recorded no interactions between these two
species at Barrow, Alaska, when they bred together in’
response to a high lemming population.
We suggest that the major relationship between the
two species is that of kleptoparasite and victim. The
mobbing reactions of the Short-eared Owls demon-
strate that they recognize Snowy Owls as potential
pirates or predators. In most records of Short-eared
Owls in the diet of Snowy Owls (e.g., Uttendorfer
1952: Mikkola 1976) it is unknown whether predation
or scavenging had occurred. The Snowy Owl has
remarkably catholic food habits (Gross 1944) and
actual predation may be of irregular or exceptional
occurrence. The rarity of documented predation and
the weakness of the Short-eared Owl’s mobbing
reaction suggest that Snowy Owl predation upon
Short-eared Owls is not significant in the biology of
either species.
414 THE CANADIAN FIELD-NATURALIST
Acknowledgments
We thank R.M. Fisher, D. P. Nugent, C. C.
Mamo, and M. Rothfels for assistance in the field, and
M. T. Myres and J. B. Gollop for their comments on
the manuscript. R. M. Fisher made his notes on
observation 6 available to us. This work was
supported by the National Research Council of
Canada and the University of Calgary.
Literature Cited
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1964-1972. Calgary Field Naturalist 3: 106-107.
Birney, E. C., W. E. Grant, and D. D. Baird. 1976. Impor-
tance of vegetative cover to cycles of Microtus popula-
tions. Ecology 57: 1043-1051.
Clark, R. J. 1975. A field study of the Short-eared Owl,
Asio flammeus (Pontoppidan), in North America. Wild-
life Monographs, Number 47.
Duffy, D. C., B. Beehler, and W. Hass. 1976. Snowy Owl
steals prey from Marsh Hawk. Auk 93: 839-840.
Gross, A. O. 1944. Food of the Snowy Owl. Auk 61: 1-18.
Hinde, R.A. 1970. Animal behaviour. 2nd_ Edition.
McGraw-Hill Book Company, New York.
Hohn, E. O. 1973. Winter hunting of Snowy Owls in farm-
land. Canadian Field-Naturalist 87: 468-469.
Keith, L. B. 1964. Territoriality among wintering Snowy
Owls. Canadian Field-Naturalist 78: 17-24.
Vol. 93
Lack, D. 1946. Competition for food in birds of prey.
Journal of Animal Ecology 15: 123-129.
Levin, S. A., J. E. Levin, and R. T. Paine. 1977. Snowy
Owl predation on Short-eared Owls. Condor 79: 395.
Lockie, J. D. 1955. The breeding habits and food of Short-
eared Owls after a vole plague. Bird Study 2: 53-69.
Meinertzhagen, R. 1959. Pirates and predators. Oliver and
Boyd, Edinburgh.
Mikkola, H. 1976. Owls killing and killed by other owls and
raptors in Europe. British Birds 69: 144-154.
Nero, R. W. 1964. Snowy Owl captures duck. Blue Jay 22:
54_55.
Pitelka, F. A., P.Q. Tomich, and G. W. Treichel. 1955.
Ecological relations of jaegers and owls as lemming
predators near Barrow, Alaska. Ecological Monographs
25: 85-117.
Salt, W.R. and J. R. Salt. 1976. The birds of Alberta.
Hurtig Publishers, Edmonton, Alberta.
Schoener, T. W. 1969. Models of optimal size for solitary
predators. American Naturalist 103: 277-313.
Tullock, R. J. 1968. Snowy Owls breeding in Shetland in
1967. British Birds 61: 119-132.
Uttendorfer, O. 1952. Neue Ergebnisse uber die Ernahrung
der Greifvogel und Eulen. Eugen Ulmer, Stuttgart.
Received 26 February 1979
Accepted 9 June 1979
—
The Biological Flora of Canada
1. Vaccinium angustifolium Ait., Sweet Lowbush Blueberry*
IVAN V. HALL,’ LEwWIs E. AALDERS,! NANCY L. NICKERSON,! and SAM P. VANDER KLOET?
'Research Station, Agriculture Canada, Kentville, Nova Scotia B4N 1J5
*Department of Biology, Acadia University, Wolfville, Nova Scotia BOP 1X0
Hall, Ivan V., Lewis E. Aalders, Nancy L. Nickerson, and Sam P. Vander Kloet. 1979. The biological flora of Canada.
1. Vaccinium angustifolium Ait., Sweet Lowbush Blueberry. Canadian Field-Naturalist 93(4): 415-430.
This contribution on Vaccinium angustifolium Ait., Sweet Lowbush Blueberry (Ericaceae), is the first of a series presenting
biological information on plants that are components of the flora of Canada. Vaccinium angustifolium is a deciduous low
shrub endemic to North America, whose range in Canada extends from the east coast of Newfoundland to Lake Winnipeg in
Manitoba. It occurs from Quebec 57°N to Virginia 38°N. To produce fruit the flowers usually require cross-pollination by
wild bees or related insects. The fruit of V. angustifolium is edible and matures during late summer. During the past 30 yr inthe
Atlantic Provinces of Canada many fields abandoned from other forms of agriculture have been brought into stands of this
species by burn-pruning and selective weed control.
Key Words: Vaccinium angustifolium, Sweet Lowbush Blueberry, biology, ecology, physiology, distribution, economic
importance.
1. Name
Vaccinium angustifolium Ait.; section Cyanococcus; Ericaceae;
V. lamarckii Camp (Camp 1943, 1945);
V. pennsylvanicum Lam. (Robinson and Fernald 1908);
V. pensilvanicum Lam. (Fernald 1950); Sweet Lowbush Blueberry;
airelle a feuilles étroites (Marie-Victorin 1964).
2. Description of the Mature Plant
(a) Raunkiaer life-form. Chamaephyte. Winter-deciduous, broad-leaved low shrub with ascending branches,
edible blue fruits, deep tap root; reproduces by seeds and rhizomes.
(6) Shoot morphology. Stems woody, average height 20 cm with maximum of 50 cm, generally glabrous, with
raised lenticels, the bark variously pigmented from yellow to deep red in autumn; buds of two types, the larger
flower buds borne terminally, the more lanceolate vegetative buds born proximally; leaves alternate ina spiral,
simple, pinnately netted, serrate glandular, elliptic or ovate-oblong, apex acute, base obtuse, the ventral surface
waxy green, the dorsal surface pale green and sometimes with light bloom; rhizomes woody bearing numerous
shoots, new growth white or pinkish (Figure la, b).
(c) Root morphology. The radicle of the seedling develops into an extensive tap root system (Hall 1957): the
root system is finely divided at the extremities and several authors state that there are no root hairs
(e.g., Addoms and Mounce 1931).
(d) Inflorescence. The members of Vaccinium section Cyanococcus are characterized by flowers borne in
racemes; members of section Vaccinium by contrast have flowers borne singly in the leaf axils. Flowers of
V. angustifolium are typically pentacyclic with five sepals, five petals fused into a bell-shaped corolla, 10
stamens in two whorls of five and fused to the corolla, and a single pistil with inferior ovary. Fruit is a true berry
bearing 10 pseudolocules each with a few to many small seeds. Bell (1957) found that the average number of
perfect seeds was 13 and of imperfect seeds was 50 per berry. Some mutants were reported by Hall, Aalders, and
Lockhart (1964). Both Vaccinium and Gaylussacia have inferior ovaries but the seeds of the former are much
smaller.
(e) Subspecies. None.
*See La Roi (1977) for notice, guidelines and schedule for contributors.
415
416 THE CANADIAN FIELD-NATURALIST Vol. 93
FIGURE |. Sweet Lowbush Blueberry Plant. A: Above, close-up of flowers: below, flowering plant. B: Above, close-up of
fruit: below, fruiting plant.
(f) Varieties and forms. Fernald (1950) recognized a puberulent var. hypolasium and a smooth-leaved var.
laevifolium House, but we do not, considering these to be part of the variation found in the species. We
recognize the following forms: V. angustifolium Ait. forma angustifolium with green leaves, fruit with a heavy
bloom, consequently a blue color, and straight stems, is the most frequent taxon (Vander Kloet 1978);
V. angustifolium forma nigrum (Wood) Boivin is characterized by blue-green leaves, fruit with little or no
bloom giving it a black color, and zigzag stems (Aalders and Hall 1963a); V. angustifolium forma leucocarpum
(Deane) Rehder is a white-fruited form.
(g) Ecotypes. Forma nigrum tends to increase more rapidly than forma angustifolium in stands that are burned
regularly (Hall et al. 1975).
(h) Chromosome numbers. Longley (1927) first reported 24 bivalent chromosomes for V. angustifolium.
Newcomer (1941) sub V. pennsylvanicum also reported that five selections had n = 24 chromosomes. Darrow et
al. (1942) are the only workers to report a diploid condition (2m = 24) but no voucher specimen for this count can
be located (Vander Kloet 1978). Counts from clones of Canadian and Maine V. angustifolium all have
2n = 48 (Hall and Aalders 1961; Bent and Vander Kloet 1976; Hersey and Vander Kloet 1976; Whitton 1964).
3. Distribution and Abundance
(a) Geographic range. Vaccinium angustifolium is a North American endemic, extending from Cape St.
Francis on the eastern tip of Newfoundland to the Pine Falls area of Lake Winnipeg in south central Manitoba
(Figure 2). It extends from 57°N in northern Quebec southward to isolated uplands in the Appalachian
Mountains of Virginia, 38°N. A complete distribution map appears in Vander Kloet (1978).
1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 417
°/
AS
800 Kilometres
Vaccinium
angustifolium Ait.
FiGuRE 2. Canadian distribution of Vaccinium angustifolium from specimens in the Department of Agriculture Herbarium,
Ottawa, Ontario (DAO): National Herbarium, National Museums of Canada, Ottawa, Ontario (CAN): and Acadia
University Herbarium, Wolfville, Nova Scotia (ACAD).
(b) Altitudinal range. Vaccinium angustifolium extends from ca. sea level to 1300 m asl in Eastern Canada to
1300-1500 m asl in Virginia. On the higher points of the Adirondack Mountains V. angustifolium is replaced
by V. boreale (Sweet Hurts) (Vander Kloet 1977).
4. Physical Habitat
(a) Climatic relations. Vaccinium angustifolium occurs ina large area of eastern North America and tolerates a
wide range of climatic conditions. The variability of this species with regard to productivity is well established
(Aalders and Hall 1963b: Hall, Aalders, and Wood 1966) but the adaptiveness of the species to different climatic
conditions requires further study. A comparison of growth under different climatic conditions was conducted
on uniform plant and soil media in 1965 and 1966 at Kentville, Nova Scotia; Normandin, Quebec; and St. John’s
West, Newfoundland. Linear shoot growth of 20 plants from seven clones at the three respective stations
averaged 183, 141, and 111 cm per plant in 1965; and 195, 153, and 70 cm in 1966. The number of flower buds
produced per plant averaged 126, 94, and 66 in 1965; and 88, 54, and 23 in 1966. Thus climatic conditions for
growth varied widely among the three stations. Vegetative growth was not notably different between 1965 and
1966 at Kentville and Normandin, but much poorer in 1966 at St. John’s West; reproductive growth was much
better in 1965 than 1966 at all three stations.
In Eastern Canada, winter temperatures for V. angustifolium and other chamaephytes are generally
ameliorated by a snow cover. Inadequate snow cover may be followed by shoot dieback to ground level in the
Lac de St. Jean area (Poirier and Dubé 1969). Unusually warm weather and salt spray from a February
hurricane, followed by cold temperatures, were probably responsible for extensive mortality of FV.
angustifolium flower buds (Figure 3).in the Fox River area of Nova Scotia in 1976. Cold winds off the
Northumberland Strait are a factor in delaying the beginning of plant growth in Inverness County, Nova Scotia
(Hall et al. 1963).
418 THE CANADIAN FIELD-NATURALIST Vol. 93
FIGURE 3. Winter injury on shoots of Vaccinium angustifolium.
In New Brunswick, Hall (1955) found that 0.5% of full sunlight under a Picea rubens — Abies balsamea
canopy (Red Spruce —- Balsam Fir) gave a minimum growth or just survival, 10% sunlight under a Berula
populifolia canopy (Oldfield Birch) gave moderate vegetative growth but no flowering, and 50% sunlight
(Openings in canopy) gave flowering and fruiting.
Seed germination occurs only after periods of prolonged rainfall in late summer or early fall. Vander Kloet
(1976a) found that seedling density was higher in a field sampled in Pictou County, Nova Scotia than in an old
field at Leeds County, Ontario and he correlated this with higher precipitation and soil moisture.
Relative humidity may play an important role in fruit production. In dry years areas along the Atlantic coast
have produced exceptional crops. Plants in the coastal fog belt are frequently covered with water droplets while
plants farther inland experience stress from lack of moisture (e.g., withered berries, browned leaf margins).
(b) Physiographic relations. One of the most important factors limiting flowering and fruit development is the
occurrence of late spring frosts on low-lying areas (Jackson et al. 1972b). Laboratory tests have shown that 6 h
of —2.2°C were detrimental and a further reduction to —3.3°C for 6 h markedly reduced fruit set (Hall, Aalders,
and Newbery 1971). Vaccinium angustifolium grows, yields, and reproduces well by rhizomes on both mineral
and organic soils provided adequate moisture and aeration are available. It tolerates a wide range of soil
conditions with best growth at low pH levels (Hall, Aalders, and Townsend 1964). In soils with much humus,
most rhizomes grow in the top 5 cm. Jackson et al. (1972a) reported that soils such as Westbrook loam and
Southampton sandy loam witha high percentage of stone or gravel provide the conditions most favorable to the
emergence, growth, and development of seedlings.
1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 419
(c) Nutrient and water relations. At the time of flower bud initiation the following suggested levels of nutrients
(as % dry weight) should exist in the leaves (Townsend and Hall 1970): N, 1.50-2.00: P, 0.08-0.12: K, 0.40-0.55:
Ca, 0.40-0.65; and Mg, 0.15-0.20. Lockhart (1959) first described the symptoms of mineral deficiency in V.
angustifolium. Rayment (1965) clearly demonstrated the beneficial effects of applications of nitrogen to stands
of V. angustifolium in Newfoundland.
Townsend et al. (1968) reported that sugars were lower, and starch was notably higher, in the rhizomes than in
the leaves. Barker et al. (1963) found that the reducing sugars in fruits of five different clones varied between 7.4
and 7.9% of fresh weight. ;
Once established, the rhizone system of this species plays an important role in preventing slope erosion. If soil
particles are washed into the network of rhizomes and shoots, new roots and shoots develop in the additional
soil, favoring the plant and retaining the soil. The taproot system (Hall 1957) may penetrate to >1 msoil depth,
allowing use of subsoil moisture reserves. Laycock (1967) states that the concentration of roots and rhizomes
near the surface in the pine barrens of New Jersey allowed V. angustifolium and other species to absorb much of
the water from light rains that fell during the growing season. The nutrient and pH requirements and
competitive ability of this polymorphic species invite further investigation.
5. Plant Communities. Within the largely undisturbed part of its range, V. angustifolium is an important
member of exposed headland vegetation, raised bogs, high moors, outcroppings on the Canadian Shield,
mountain summits in the Gaspé, as well as in the herb—dwarf shrub stratum of open to moderately shaded pine
to spruce woodlots and forests on coarse textured, mainly infertile soils in the Acadian, Great Lakes — St.
Lawrence and Boreal Forest Regions of Rowe (1972), from Lake Winnipeg to the Atlantic coast.
But the species is most abundant in disturbance communities which result from clear-cutting, forest fires, and
abandonment of agricultural land. The latter is.especially noteworthy in the Maritime Provinces where, witha
decline in soil fertility, land was abandoned during the early part of this century, providing many old field
habitats for this species to colonize. Conners (1972) reports that by the late 1940s several of these fields were
being cultivated. With the introduction of frozen foods on the American market a rapid development of the
stands and industry took place.
Table | shows the contrasts among the floristic composition of three community types: (1) blueberry barrens
on mainland Nova Scotia, (2) granite outcroppings near Kaladar, Ontario and, (3) the boreal conifer-hardwood
forests of the Great Lakes region. Aside from V. angustifolium, the three community types have floristically
littie in common. Out of 71 species, excluding lichens and those with less than five occurrences, only V.
myrtilloides (Sour-top Blueberry), Gaultheria procumbens (Teaberry), Fragaria virginiana (Wild Strawberry),
Rumex acetosella (Sheep Sorel), and Maianthemum canadense (Wild Lily-of-the-Valley) are shared.
Maycock and Curtis (1960) have shown that V. angustifolium is an important component in both dry and wet
sites of the boreal coniferous forest, but is unimportant on mesic sites. In Wisconsin, Curtis (1959) showed that it
is the most prevalent groundlayer species in the northern dry forest and also an important constituent of
northern dry-mesic forest, northern wet forest, bracken-grassland, and open bog. Lamoureux and Grandtner
(1977) showed that this species was important in dune formation on Iles-de-la-Madeleine. Lavoie (1968) has
described the Jack Pine ( Pinus banksiana) forests witha Kalmia angustifolia-Vaccinium understory for the Lac
de St. Jean area of Quebec. Community descriptions and tabulations of V. angustifolium in the Maritime.
Provinces are given by Hall (1955, 1959, 1975), Hall and Aalders (1968), and Hall et al. (1973, 1974, 1976).
6. Growth and Development
(a) Morphology. In the lowbush blueberry seedling (Figure 4) the oblong-elliptical cotyledons are about 2 mm
long. The first seedling leaves are much smaller and usually more elliptical than mature leaves, making
identification of young seedlings difficult. Stages in seedling development are given by Eaton and Hall (1961).
Plants rarely flower or produce rhizomes until 4 yr after germination (Hall 1953). After plants reach a total
width of 30 cm and rhizome growth occurs in several directions, expansion of the clone is more rapid. Rhizomes
may grow up to 10 cm/yr on mineral soils and up to 50 cm on organic soils. Clones with intact rhizomes 10 m
long have been excavated in Kings County, Nova Scotia. The minimum age of a parent clone may be estimated
by counting growth rings from rhizome cross-sections (Figure 5). The importance of rhizome growth in the
expansion of the clone has been outlined by Barker and Collins (1963b).
(b) Physiology. Forsyth and Hall(1965) have shown that photosynthesis is more rapid in the early morning and
that temperature, CO, and leaf age affect photosynthetic rates. Rates of apparent photosynthesis in shoots at
different light intensities have been reported by Bonn et al. (1969). Red leaf infected leaves (see 9(c)) and genetic
mutant leaves had abnormally lower photosynthetic rates (Hall, Forsyth, Lockhart, and Aalders 1966). Strong
420 THE CANADIAN FIELD-NATURALIST Vol. 93
TABLE 1— Vaccinium angustifolium Aiton, associated species on six granitic outcroppings from Kaladar, Lennox-Addington
County to Mount Fitzsimmons, Leeds County, eastern Ontario: two barrens* on mainland Nova Scotia: and I! dry Boreal
Conifer — Hardwood Forests of the Great Lakes Region**
Frequency of occurrence (%)
Eastern Mainland Great Lakes
Species Ontario Nova Scotia region
Bare ground, rock, or litter 789 (21.6) 1716 (17.2) NA
Epipetric lichens 522 (14.3) — NA
Fruticose lichens PON GSES) —- NA
Polytrichum commune 28 ( 0.8) 199 ( 2.0) NA
Polytrichum jJuniperinum ISU ( 41) — NA
Polytrichum piliferum 33 ( 0.9) = NA
Lycopodium annotinum*** — = (12)
Lycopodium clavatum _ 6 (21)
Lycopodium obscurum a 44 ( 0.4) (22)
Lycopodium complanatum — 6 ( 0.06) ( 1)
Pteridium aquilinum _ 59 ( 0.6) (78)
Juniperus communis @il ( 25) — —
Danthonia spicata 159 ( 4.4) 536 ( 5.4) =
Deschampsia flexuosa 144 ( 3.9) = ==
Poa compressa 30 ( 0.8) —- —
Agrostis scabra 6 ( 0.2) — —
Panicum subvillosum = 14 ( 0.1)
Oryzopsis asperifolia — — ( 8)
Carex pensylvanica 228 ( 6.2) —
Carex nigromarginata 23 ( 0.6) — =
Carex umbellata 8 ( 0.2) = =
Luzula multiflora = 6 ( 0.06) a
Clintonia borealis — — (17)
Maianthemum canadense 15 ( 0.4) 8 ( 0.08) (66)
Salix bebbiana — 335) ((_ )315))) =
Betula populifolia _— 30 ( 0.3) —
Corylus cornuta — = (23)
Rumex acetosella AV) (( 1.3) 4( 0.04) (ey)
Comandra umbellata 66 ( 1.8) i
Anemone quinquefolia = a
Dalibarda repens — —
Spiraea alba 18 ( 0.5) —
Spiraea latifolia _ 60 ( 0.6) —
Spiraea tomentosa — 29 ( 0.3)
Amelanchier sanguinea ihe @.33)) — —
Amelanchier spicata 19 ( 0.5) — —
Prunus virginiana 9 ( 0.3) — —-
Prunus serotina 7 (( O22) — —
Pyrus melanocarpa 163 ( 4.5) -- a
Rubus arundelanus 12 ( 0.3) — =
Rubus hispidus = 41 ( 0.4) —
Rubus idaeus -- — ( 6)
Waldsteinia fragarioides — — (18)
Fragaria virginiana 5) (( W310) 39 ( 0.4) (#5)
Potentilla tridentata — 46 ( 0.5) =
Potentilla simplex = 84 ( 0.8) —
Polygala paucifolia _ — (22)
Rhus copallina 9 ( 0.2) — —
Rhus typhina 11 ( 0.3) = —
Aralia nudicaulis — — (33)
Cornus canadensis — 389 ( 3.9) (61)
Ledum groenlandicum _- 14 ( 0.1) —
Rhododendron canadense — 106 ( 1.1) —
1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 421
TABLE |—(Concluded)
Frequency of occurrence (%)
Eastern Mainland Great Lakes
Species Ontario Nova Scotia region
Kalmia angustifolia — STM (B21) —
Arctostaphylos uva-ursi Ol @ ey) — —
Gaultheria procumbens 23 ( 0.6) 348 ( 3.5) (21)
Vaccinium myrtilloides 32 ( 0.9) 1 ( 0.01) (20)
Vaccinium angustifolium 528 (14.5) 2139 (21.4) (51)
Vaccinium vitis-idaea — Po (O27) —
Gaylussacia baccata sy) (IS) = =
Trientalis borealis — — (39)
Melampyrum lineare — — (19)
Linnaea borealis = — (12)
Viburnum rafinesquianum 21 ( 0.6) — =
Lonicera canadensis — 6 ( 0.06) =
Diervilla lonicera 8:((022) = ==
Helianthus divaricatus 26 ( 0.7) = =
Aster cordifolius 8 ( 0.2) — —
Aster macrophyllus — = (33)
Solidago puberula - 648 ( 6.5)
Solidago rugosa = 6 ( 0.06) =
Solidago graminifolia — 15 ( 0.15) ( 3)
Antennaria neglecta = 120812) (a)
Prenanthes trifoliolata = 5 ( 0.05) —
Hieracium aurantiacum = 8 ( 0.08) (12)
Hieracium pratense — 5 ( 0.05) —
34 spp. with fewer than 5 occurrences 46 ( 1.3) 10 ( 0.1) NA
Totals 3650 (100) 10,000 (100) 11 Stands
*Data from Hall and Aalders (1968) slightly modified and reduced.
**Data from Maycock and Curtis (1960) slightly modified and reduced.
***Scientific nomenclature of vascular plants follows Gray’s Manual of Botany (Fernald 1950).
vegetative growth results when plants of different provenances are given 16 h light and 8 h dark periods per 24 h
at 18°C. Flower buds are formed when plants are given 12 h light and 12 h dark per 24 h at 18°C. Hall et al.
(1970) have shown that leaf anthocyanin concentration increases as the temperature decreases. Vaccinium
angustifolium leaves develop brilliant red or yellow coloration in autumn.
(c) Phenology. Vaccinium angustifolium overwinters in a leafless state, with twigs yellow to reddish brown. At
Kentville, vegetative and flower buds swell in early May if air temperatures have exceeded 10°C for 3-4 d.
Flowering occurs from late May to mid-June. In cooler coastal areas, flowering may be delayed 2-3 wk (Bell
1953). Leaf development precedes, is concomitant with, or follows flowering depending on the particular clone.
Leaves harden by mid-July, turn red in late August, and absciss by late October. In seasons following a burn-
prune, vegetative growth commences a week after flower bud expansion and terminates in early July.
Termination of shoot growth as evidenced by a black tip in the apical meristem (Bell 1950) occurs earlier on
older unpruned bushes (Barker and Collins 1963a). Berries ripen in early August at Kentville, 2-3 wk earlier
than in Cumberland County, Nova Scotia (Aalders et al. 1972). Flower primordia begin to develop shortly after
cessation of vegetative growth (Bell and Burchill 1955a), but continue until late October if air temperatures
remain >0°C with extended periods greater than 10°C. Thus temperature is an important regulator of plant
phenology. But other factors including day length are important (Hall et al. 1970). At Lac de St. Jean, Quebec,
flower bud formation is poor in some years, probably owing to early frosts. Halland Ludwig (1961) have shown
that different clones react differently to day length and temperature. By winter, primordia ofall the floral organs
are microscopically recognizable (Bell and Burchill 1955b).
Kender (1968) found that growth potential of rhizome buds was greater in early spring and late summer than
in July.
422 THE CANADIAN FIELD-NATURALIST Vol. 93
FIGURE 4. Sweet Lowbush Blueberry seedlings. Approximate age 3 yr upper left and 4 yr lower right. The latter has begun to
form a rhizome (indicated by arrow).
7. Reproduction
(a) Floral biology. The flowers are insect pollinated, but not very attractive to bees, probably because the nectar
volatiles are simple acetaldehyde and ethanol (Hall, Forsyth, Lightfoot, and Boch 1971). The stamens and pistil
are functional as soon as the flower fully opens. The pistil remains highly receptive under field conditions for4 d
but fertility drops to about 20% by the 7th d (Wood 1962). Wood (1961a) also found that nectar volume and
weight of nectar sugar increased with the flower age. After pollination the corolla turns pink and deteriorates
very rapidly. This is associated with a marked increase in ethylene production (Hall and Forsyth 1967) anda
high rate of respiration (Forsyth and Hall 1969) which persists through the small green fruit stage.
Glasshouse studies indicate that flowers selfed using pollen from the same flower or any flower of the same
clone do not generally set fruit (Aalders and Hall 1961). The results of intercrossing and selfing six clones of V.
angustifolium are given in Table 2. The vascular anatomy of the ovary has been described by Bell and Giffin
(1957).
(b) Seed production and dispersal. It is rare to find a seedless fruit of V. angustifolium even in highly productive
clones. Table 2 shows the number of seeds per berry in crosses completed in 1977 and 1978 among six clones.
1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 423
Seeds are spread in the droppings of birds and mammals. The American Robin (Turdus migratorius) isa major
seed disperser in southwestern New Brunswick as it often feeds and migrates just before fruit harvest by man
(Eaton 1957). We have found seeds of V. angustifolium in the droppings of Black Bear ( Ursus americanus), Red
Fox (Vulpes vulpes), and Raccoon (Procyon lotor).
TABLE 2—Cross- and self-fertility of six selected clones of Vaccinium angustifolium in glasshouse trials, Kentville,
Nova Scotia, 1977-1978
Percent of
pollinated No. seeds Percent
flowers forming No. seeds per 100 seed
Female Male fruits! per berry? pollinations germinations3
Augusta ™X Brunswick 100 31 3100 88
x Chignecto 70 24 1680 97
x 510 65 25 1625 98
xX ME3 81 55 4485 93
x ME4161 61 49 3026 88
x Self 56 10 568 89
Brunswick X Augusta 0 — 0 =
x Chignecto 0 = 0 --
x 510 2 16 30) 94
x ME3 2 20 49 90
x ME4161 0 os 0 —
x Self 0 — 0 =
Chignecto X Augusta 74 30 2249 88
x Brunswick 65 17 1131 91
x 510 5) 24 1298 90
xX ME3 56 28 1548 98
x ME4161 45 25 1125 95
xX Self 0 — 0 =
510 x Augusta 12 46 580 84
x Brunswick 95 34 3230 80
x Chignecto 80 33 2632 86
x ME3 1] 4] 461 82
x ME4161 0 oo 0 =
x Self 0 — 0 =
ME3 x Augusta 18 37 644 89
x Brunswick ~ 65 39 2535 94
x Chignecto 14 38 521 89
x 510 6 22 140 95
x ME4161 l 14 18 93
xX Self 0 = ) —
ME4161 X Augusta 42 21 892 97
x Brunswick 29 23 673 93
x Chignecto 26 16 410 90
x 510 39 23 907 91
<x ME3 75 30 2235 92
x Self 14 6 77 74
Crosses 30.6% 29 1241 82
Selfs 11.7% 8 108 90
'Based on a sample of 20-80 hand-pollinated flowers.
2Seeds used here were the perfect ones. according to Bell (1957).
3Based on four lots of 50 seeds each, or on fewer if 200 not available.
424 THE CANADIAN FIELD-NATURALIST Vol. 93
HEART WOOD
PRIMARY XYLEM
BARK
FIGURE 5. Cross-section of a Vaccinium angustifolium rhizome showing 21 growth rings. Counting these rings provides an
estimate of the minimum age of the parent clone.
The number of seed (up to 64) found ina berry depends on genetic factors and environmental conditions. If
weather is favorable (warm and dry) for insect pollination during the period of stigma receptivity, berries will
have higher seed counts.
(c) Seed viability and germination. Fresh seed of V. angustifolium extracted from well ripened berries
germinates readily at 21°C under 16 h light per 24-h period (Table 2) and usually takes 21 d. Germination is
greater in light than in dark (G. W. Wood, personal communication). Seeds can be stored dry or frozen in fruit
for 6 mo with no major reduction of viability (Aalders and Hall 1975). Seeds of V. angustifolium germinate well
when sown on a soil mix, pH 4.7 in the glasshouse.
(d) Vegetative reproduction. Asexual reproduction occurs when the rhizomes are cut or killed by fire, shading,
burrowing and/ or frost action whereas sexual reproduction predominates in areas where the soil has recently
been denuded by cultivation, flooding, or blow down of forest trees.
Plants may be propagated from either rhizome or stem cuttings. At Kentville, stem cuttings are taken just
after black tip formation (6(c)) in early July and rooted under intermittent mist for 6 wk, with about 80% success
(Hall et al. 1977). Studies of in vitro culture techniques for propagation are in progress (Nickerson and Hall
1976: Nickerson 1978).
8. Population Structure and Dynamics
Since dispersal is through bird and mammal droppings, one would expect a clumped dispersion pattern, and
random sampling of three abandoned meadows with 28 |-m? quadrats in Pictou County, Nova Scotia gave a
mean + SE of |.1 £0.39 seedlings, variance of 4.54, and a variance mean ratio of 4.3:1, confirming that
expectation. One quadrat had a clump of 17 seedlings, the remainder one or none, rarely two seedlings.
Seedlings may initially be clumped but as Vander Kloet (1976a) has shown, both in Nova Scotia and Ontario,
seedling mortality is very high, 1.e., > 99% of germinating seeds died even under partially controlled conditions.
1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 425
The shoot crown diameters of plants established by seed can be converted to age classes according to a
formula devised by Eaton and Hall (1961). Using this size-age relationship, Vander Kloet (1976b) found a
marked difference in the age structure of V. angustifolium in old fields in eastern Ontario and eastern Nova
Scotia. In the former, all seed-established plants exceeded 40 yr of age: in the latter the 11-20 yr age class was
the most frequent.
On stable habitats such as mountain summits, exposed headlands, or outcroppings in the Canadian Shield,
colonies of V. angustifolium may attain considerable age. Clones attain large size (>10 m diameter) and age
(150 yr). Vander Kloet (unpublished data) studied the species composition and population patterns of
granitic outcrop communities in the Thousand Island region of Ontario and New York over a 10-yr period.
Little or no change in species composition or abundance was observed; however, the positions of colonies of V.
angustifolium, Juniperus communis (Common Juniper), and Pyrus melanocarpa (Black Chokeberry) shifted
laterally. Vigorous peripheral branches of Juniperus shaded out colonies of Vaccinium and Pyrus, while the
latter two invaded the senescing centers of Juniperus colonies. Similarly Pyrus and Vaccinium invaded the
senescent portions of each other’s colonies.
Turnover rates in V. angustifolium populations may be more rapid in the maritime provinces where the
species is often a common component of old field succession, reaching peak abundance 15-25 yr after
abandonment. By the judicious use of fire, this seral stage can be maintained as blueberry barrens for many
years. Ordinarily the low growing V. angustifolium is soon shaded out by a variety of hardwood or conifer tree
species. Once the forest canopy fully develops, it becomes uncommon and/or sterile. Vaccinium myrtilloides
has a much higher survival potential than V. angustifolium in the mature Acadian boreal forest (Hall 1959).
9. Interaction with Other Species
(a) Competition. Competing plant species may be treated in four ecological site groups: (1) undisturbed
natural; (2) post-logging; (3) old field; (4) blueberry crop.
On undisturbed natural sites within its range of suitable habitats, V. angustifolium has many competitors, as
may be seen in Table 1. In relatively stable and oligotrophic Pinus banksiana woodlands and Picea mariana
(Black Spruce) forests of the Boreal Forest and Great Lakes — St. Lawrence Regions (Rowe 1972) east of Lake
Winnipeg, its major competitors for nutrients, water, and light in the herb-dwarf shrub stratum include I.
myrtilloides, Cornus canadensis (Bunch Berry), Gaultheria hispidula (Creeping Snowberry), Coptis
groenlandica (Goldthread), Pteridium aquilinum (Bracken Fern), Clintonia borealis (Corn-lily), Comptonia
peregrina (Sweet-fern), Maianthemum canadense, and Lycopodium spp. Major competitors in the low shrub
stratum include Kalmia angustifolia (Lambkill), Ledum groenlandicum (Labrador-tea), and Diervilla
lonicera (Bush Honeysuckle).
On post-logging sites in eastern Canada, surviving and/or invading V. angustifolium populations must
compete with surviving species that respond favorably to clearing, e.g., Pteridium aquilinum (Cody and
Crompton 1975), Dennstaedtia punctilobula (Hay-scented Fern) (Cody et al. 1977), and Cornus canadensis
(Hall and Sibley 1976), as well as with aggressive invader species, e.g., Epilobium angustifolium (Fireweed).
On old field sites in the Acadian Forest Region (Rowe 1972) V. angustifolium is an important seral species in
the transition stage between field and forest. On coarse-textured soils in old fields of Prince Edward Island,
Myrica pensylvanica (Bayberry) succeeds V. angustifolium before the Picea glauca excludes both (Hall 1975).
On light sandy soils of the Lac de St. Jean of Quebec, Comptonia peregrina is a major invader of blueberry
stands (Lavoie 1968).
Members of the blueberry crop group offer competition because they thrive under the cultural practices
pertaining to lowbush blueberry culture. Probably the strongest competitor for space is Kalmia angustifolia
(Hall et al. 1973). Its stem growth following burn-pruning exceeds that of V. angustifolium (Hall and Aalders
1968). Fertilizing stands of lowbush blueberry increases the growth of Pyrus melanocarpa relative to that of FV.
angustifolium (Hall et al. 1978). On poorly drained areas Rhododendron canadense (Rhodora) and Spiraea
latifolia (Meadow-Sweet) (Hall et al. 1974) replace V. angustifolium.
(b) Symbiosis. The principal native pollinators in Maine and eastern Canada are species of Halictidae and
Andrenidae although a few species of Bombidae, Anthophoridae, Colletidae, and Xylocopidae are of some
importance (Boulanger et al. 1967). In eastern Ontario Vander Kloet (1976c) found that the solitary bees
Andrena vicina and A. carlini, and the bumblebees Bombus bimaculatus, B. terricola, and B. ternarius were the
important oe Wood (1961b) found honeybees effective during a short period of bloom in New
Brunswick.
Mycorrhizal associations in V. angustifolium were described by M. MacArthur (1955. Mycorrhiza in the
426 THE CANADIAN FIELD-NATURALIST Vol. 93
blueberry. /n Horticulture Division, Central Experimental Farm, Ottawa, Progress Report 1949-53. pp. 71-72)
but the fungi involved were not identified.
(c) Predation and parasitism. Foliage of V. angustifolium is eaten by Black Bear ( Ursus americanus), Eastern
Cottontail (Sy/vilagus floridanus), and White-tailed Deer (Odocoileus virginianus). Fruits are eaten by a
number of mammals and many birds (Martin et al. 1951).
The important insects are Blueberry Maggot, Rhagoletis mandax Cn.; Black Army Cutworm, Actebia
fennica (Yausch.); Chainspotted Geometer, Cingilia catenaria (Drury); Blueberry Flea Beetle, Altica sylvia
Mall.: Blueberry Casebeetle, Chamisus cribripennis (LeConte) (see Wood 1978): Blueberry thrips, Frankliniella
vaccinni Morgan and Catinathrips kainos O'Neill: Blueberry Tipworm, Contarinia vaccinii Felt.: sawflies,
Neopareophora litura Klug, Pristiphora idiota Nort., Pristiophora sp.; Red-striped Fireworm, Aroga
trialbamaculella Chamb.; and Stem Galler, Hemadas nubilipennis Ashm.
The stage of the life cycle of these insects which affects V. angustifolium, the manner of infection, the
symptoms for recognition, and other details are given in Hall et al. (1975).
The important fungal diseases (Conners 1967) are monilinia blight caused by Monilinia vaccinii-corymbosi
(Reade) Honey: botrytis blight (Borrvtis cinerea Pers.): red leaf (Exobasidium vaccinii Wor.); witches-broom
(Pucciniastrum goeppertianum (Kuhn) Kleb.); dieback (Diaporthe vaccinii Shear); powdery mildew
(Microsphaera penicillata (Wallr. ex Fr.) Lev. var. vaccinii (Schw.) W. B. Cke.): leaf rust (Pucciniastrum
vaccinii (Wint.) Jorstad); leaf spot (Septoria sp.); Gloecsporium leaf spot (G/oeosporium minus Shear); and
canker (Godronia cassandrae Pk. f. vaccinii Groves). A V. angustifolium plant which showed symptoms of
shoestring virus disease was reported by Lockhart and Hall (1962).
(d) Toxicity and allelopathy. None reported to date.
10. Evolution and Migration. Camp (1942), on the basis of meager and unsubstantiated evidence (Vander
Kloet 1978), assumed that V. angustifolium was a diploid species (1.e., gametes had 12 chromosomes) but as was
pointed out in section I(/) this cannot be confirmed. Therefore his allopatric speciation model for the species is
also doubtful since Camp (1942) argued, rightly, that processes leading up to speciation in diploid populations
differ markedly from those which give rise to tetraploid species. Vander Kloet (1977) has postulated a recent
hybrid origin for V. angustifolium. He proposes that V. boreale migrated south or moved down mountain
slopes where it came into contact with V. pallidum Ait. (Upland Low Blueberry) of oak-pine woods. Both of
these species are diploids and by spontaneous chromosome doubling in a hybrid, an allotetraploid species is
plausible.
Probably V. angustifolium migrated into the northern part of its habitat from the southern United States
following the retreat of the last glacial ice. Birds doubtless accelerated this migration by dispersing seeds across
water bodies and other habitat barriers.
An initial attempt has been made to separate clones of V. angustifolium on the basis of chlorophyll and
anthocyanin content of bark from shoots (Wood and Barker 1963).
11. Response Behavior
(a) Fire. In natural communities or managed forests V. angustifolium survives wild fire or controlled burning
below ground. Recolonization occurs by rhizome sprouting. Commercial stands are burn-pruned every second
year resulting generally in unbranched stems which have more flower buds per stem and more flowers per bud
than on older wood. There were no significant differences between fall- and spring-burned plants with respect to
amount of shoot growthand number of flower buds per shoot (Hall, unpublished). Burning after the plants were
in full leaf was detrimental to new shoot growth and flower bud formation (Eatonand White 1960). Black (1963)
found that total fruit production over a 9-yr period was greater from burning every second year than from
burning every third year. Smith and Hilton (1971) found that improved lowbush blueberry performance in
Ontario after burning resulted mainly from the stimulative effects of nutrients in ash deposited on the surface
soil.
(b) Grazing and harvesting. In New Brunswick, sheep were observed to graze grasses and sedges selectively
rather than feed on V. angustifolium and V. myrtilloides (Hall, I. V. 1954. Ecological studies. Ja Dominion
blueberry substation, Tower Hill, New Brunswick progress report 1949-1953. Canada Department of
Agriculture. pp. 18-23). Removal of shoot tips by browsing White-tailed Deer (Odocoileus virginianus) results
in lateral branching.
1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 427
(c) Flooding. During the dormant period V. angustifolium can withstand considerable flooding such as that
which occurs in many bogs, but it is not characteristic of wet habitats such as marshes or lake margins.
(d) Drought. During prolonged drought in early summer shoot growth is reduced. Dry weather later in the
season results in shriveling of fruit and reduced flower bud formation. Irrigation prior to harvest substantially
increases fruit size.
(e) Herbicides. 2,4-D causes a twisting of the terminal growth followed by browning and leaf fall. No data are
available on other herbicides.
12. Relationship to Man
The fresh fruits of V. angustifolium have been a part of man’s diet in North America since prehistoric times.
Indians dried and pulverized fruits for blending with meat (Hedrick 1919). European settlers collected and
preserved the fruit for jam, jelly, and preserves as well as eating them raw. These are still the main uses, but new
products such as muffin mixes, ice cream, yogurt, and wine (Hope 1965) are using considerable quantities of
fruit. For commercial use the fruits are quick-frozen by passing them through a tunnel of air at -29° to -34°C.
For many years the only fruit markets for the species were in Canada and the United States. Recently the fruit
has found some acceptance in western Europe where it competes with the European V. myrtillus L.
The area of lowbush blueberries under management is expanding each year in the Atlantic Provinces, which
have programs for assisting growers to develop existing stands of V. angustifolium. Agriculture Canada is also
providing funds through the New Crop Development Fund to determine the cost and feasibility of establishing
plantings of this species within its range.
Acknowledgments
The authors acknowledge the technical assistance of A. T. Lightfoot, A. C. Brydon, M. A. Spicer, and B. S.
Brooks.
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Vander Kloet, S. P. 1976a. A comparison of the dispersal and seedling establishment of Vaccinium angustifolium (the
lowbush blueberry) in Leeds County, Ontario and Pictou County, Nova Scotia. Canadian Field-Naturalist 90: 176-180.
Vander Kloet, S. P. 1976b. A novel approach to sampling Vaccinium populations. Canadian Journal of Botany 54: 669-671.
430 THE CANADIAN FIELD-NATURALIST Vol. 93
Vander Kloet, S. P. 1976c. Nomenclature, taxonomy, and biosystematics of Vaccinium section Cyanococcus (the blue-
berries) in North America. |. Natural barriers to gene exchange between Vaccinium angustifolium Ait. and Vaccinium
corymbosum L. Rhodora 78: 503-515.
Vander Kloet, S . P. 1977. The taxonomic status of Vaccinium boreale. Canadian Journal of Botany 55: 281-288.
Vander Kloet, S. P. 1978. Systematics, distribution, and nomenclature of the polymorphic Vaccinium angustifolium Aiton.
Rhodora 80: 358-376.
Whitton, L. 1964. The cytotaxonomic status of Vaccinium angustifolium Aiton in commercial blueberry fields of Maine.
Ph.D thesis, Cornell University, Ithaca, New York. 150 pp.
Wood, F. A. and W. G. Barker. 1963. Stem pigmentation in lowbush blueberry. Plant Physiology 38: 191-193.
Wood, G. W. 196la. The association between age of inflorescence and nectar production in the lowbush blueberry
Vaccinium angustifolium. Canadian Journal of Botany. 39: 1037-1040.
Wood, G. W. 1961b. The influence of honeybee pollination on fruit set of the lowbush blueberry. Canadian Journal of Plant
Science 41: 332-335.
Wood, G. W. 1962. The period of receptivity in flowers of the lowbush blueberry. Canadian Journal of Botany 40: 685-686.
Wood, G. W. 1978. Studies on Neochlamisus cribripennis (Le Conte) with particular reference to its reproductive biology.
PhD. thesis, University of New Brunswick, Fredericton, New Brunswick. 95 pp.
Received 5 June 1978
Accepted 17 April 1979
Notes
Mosses New to Ontario and Quebec
ROBERT R. IRELAND! and GILDA BELLOLIO-TRUCCO?
‘National Museum of Natural Sciences, National Museums of Canada, Ottawa, Ontario KIA 0M8
239 Commanche Drive, Ottawa, Ontario K2E 6E8
Ireland, Robert R. and Gilda Bellolio-Trucco. 1979. Mosses new to Ontario and Quebec. Canadian Field-Naturalist
93(4): 431-433.
Five mosses are reported new to the bryoflora of Ontario (Cinclidium latifolium, Drepanocladus revolvens var. intermedius,
Hypnum hamulosum, Myurella tenerrima, Orthothecium strictum) and five new to Quebec (Atrichum tenellum, Fissidens
obtusifolius, Hypnum hamulosum, Orthotrichum strangulatum fo. lescurii, Pseudoleskeella catenulata).
Key Words: mosses, Ontario and Quebec flora, geographical distribution, new records.
Recent collecting by the authors and others have
accounted for several new additions to the moss flora
of Ontario and Quebec. The Ontario records are new
to the checklist of Ireland and Cain (1975), while the
Quebec mosses are not listed by Lepage (1947),
Kucyniak (1948, 1949, 1950a, b, c, 1952a, b, c, 1961),
Love et al. (1958), LeBlanc (1949, 1951, 1954, 1963),
or Masson (1967). In the case of Fissidens obtusifolius
Wils., Lepage (1947) reported it for Quebec but
LeBlanc (1963) later deleted it from the flora.
The purpose of this paper is to report four species
and one variety new to Ontario and four species and
one form new to Quebec. Voucher specimens are
deposited in the National Herbarium of Canada
(CANM) in Ottawa.
New to Ontario
Cinclidium latifolium Lindb.
Kenora District: 56°45’N, 88°45’W, coastal raised
beach-sedge meadow system, Kershaw, July 1972
(CANM 242656).
An interesting moss collected by Kershaw (1974) in
his studies of the raised beaches in northwestern
Ontario, this species is a circumpolar arctic-alpine
calciphile that was previously known in Canada from
northern Manitoba, Yukon Territory, and Northwest
Territories. Mogensen (1973) has mapped its world-
wide distribution.
Drepanocladus revolvens var. intermedius (Lindb. ex
C. Hartm.) Grout
Bruce County: Cemetery Road, Caesar’s Bog, C.
Williams 1107. Cochrane District: island in Moose
R. near mouth of Abitibi R., 51°03’N, 80°57’W,
Baldwin B-22. Rainy River District: 5 mi [8 km]
NE of Gameland, Garton 9263. Thunder Bay
District: Sibley Peninsula, bog at NW corner of
Middlebrun Bay, 2.75 ml [4.4 km] NE of Silver
Islet, Garton 4362.
431
Ireland and Cain (1975) considered this variety a
synonym of the typical variety; however, there has
been so much debate (Steere 1978) regarding its
taxonomic status that we have decided to recognize it
as a distinct taxon until a detailed study can be made.
The variety intermedius is a calciphile that is also
known from British Columbia, Newfoundland,
Yukon Territory, and Northwest Territories.
Hypnum hamulosum B.S.G.
Kenora District: 56°45’N, 88°45’W, coastal raised
beach-sedge meadow system, Kershaw, July 1972
(CANM 242653). Verified by H. A. Crum.
Another calcicolous arctic-alpine species, like Cin-
clidium latifolium, that was collected by Kershaw
(1974) in his raised beaches studies, this species has
been reported for all Canadian provinces and terri-
tories except Prince Edward Island and New Bruns-
wick. Its presence in Newfoundland (Macoun and
Kindberg 1892) and Nova Scotia (Erskine 1968) seems
doubtful, however. Ando (1972) has mapped the
worldwide distribution of this Hypnum, as well as
others that occur in North America. .
Myurella tenerrima (Brid.) Lindb.
Thunder Bay District: Ourmet Canyon, 10 km W of
Dorion, 48° 43’N, 88°40’W, Garton 11634, 15365,
15371; canyon at NW corner of Cavern Lake,
16 km NNW of Dorion, 48°50’N, 88°41’W, Garton
14997, 15098.
This calcicolous arctic-alpine species is now known
from all provinces and territories in Canada except
Saskatchewan, Newfoundland, New Brunswick, Nova
Scotia, and Prince Edward Island. Its occurence in
Ouimet and Canyon Lake canyons is extremely
interesting as other arctic mosses, such as Au/acomni-
um acuminatum (Lindb. & H. Arnell) Kindb. (Wil-
liams 1968) and A. turgidum (Wahlenb.) Schwaegr.
also occur there, reaching their southern limit in these
two Ontario localities. These mosses are arctic relicts
432
that have been able to survive at the bottom of these
canyons far south of their present ranges because of
the cold microenvironment created among huge
boulders.
Orthothecium strictum Lor.
Kenora District: south shore Attawapiskat River,
52°52’N, 83°45’W, Riley 6512.
This is a circumboreal arctic-alpine calciphile that is
also known in Alberta, Quebec, Newfoundland,
Yukon Territory, and Northwest Territories.
New to Quebec
Atrichum tenellum (Roehl.) B.S.G.
Gatineau County: Gatineau Park, trail N of Cham-
plain Lookout, NW of Hull, 45°30’N, 75°54’W,
Treland, Iwatsuki & Kuc 9684. Determined by G. L.
Smith.
This collection was originally identified as A.
crispum (James) Sull. & Lesq. but G.L. Smith
discovered the misidentification and informed the
senior author (personal communication) that it
represented a species new to North America. Ireland
(1971) subsequently reported the species for southern
British Columbia and Quebec but without specific
collection data. The species is now known from
scattered localities in British Columbia, Manitoba,
Ontario, Labrador, Newfoundland, Prince Edward
Island, and New Brunswick.
Fissidens obtusifolius Wils.
Gatineau County: Stag Creek, near Low, ca.
45°48’N, 76°07’W, Ireland & Bellolio-Trucco
18099.
This distinct calciphile, which was reported by
Lepage (1947) for Quebec, was later deleted from the
flora by LeBlanc (1963). It is probably rare in the
province but it may eventually be found in other
localities in southern Quebec. It seems to be restricted
to eastern Canada where it is also known from
southern Ontario.
Hypnum hamulosum B.S.G.
James Bay region, 54°53’N, 79°08’W, Lethiecq
OFB-E 11498.
This species is also reported as new to Ontario in
this paper.
Orthotrichum strangulatum fo. lescurii (Aust.) Vitt
Gatineau County: Gatineau Park, Luskville Falls,
NE of Luskville, 45°32’N, 76°00’W, Ireland &
Bellolio-Trucco 17922. Determined by D. H. Vitt.
This form differs from the typical form strangu-
latum, which is also known from Quebec, in several
minor variations of the gametophyte and sporophyte
(Vitt 1973). The form /escurii has been reported for
British Columbia and Ontario (Grout 1935) but its
occurrence in British Columbia is doubtful according
to Vitt (personal communication).
THE CANADIAN FIELD-NATURALIST
Vol. 93
Pseudoleskeella catenulata (Brid. ex Schrad.) Kindb.
Gatineau County: Paugan Falls, 1 mi[1.6 km] E of
Low, ca. 45°49’N, 75°5S’W, Ireland & Bellolio-
Trucco 18050.
This calcareous species is also known from Ontario,
Labrador, and Northwest Territories. Lewinsky
(1974), who mapped the worldwide distribution,
showed a record in the Gaspé Peninsula but we have
seen neither a literature report nor a herbarium
specimen for that locality.
Acknowledgments
We thank H. A. Crum, University of Michigan,
G. L. Smith, New York Botanical Garden, and D. H.
Vitt, University of Alberta, for identifying or verifying
some of the collections.
Literature Cited
Ando, H. 1972. Distribution and speciation in the genus
Hypnum in the circum-Pacific region. Journal of the
Hattori Botanical Laboratory 35: 68-98.
Erskine, J. S. 1968. An introductory moss flora of Nova
Scotia. Nova Scotia Museum, Occasional Paper 6,
Science Series 4. 110 pp.
Grout, A. J. 1935. Moss flora of North America, north of
Mexico. Newfane Volume 2(2). pp. 67-138.
Ireland, R. R. 1971. /n Moss flora of the Pacific Northwest.
By E. Lawton. Hattori Botanical Laboratory, Nichinan.
362 pp.
Ireland, R.R. and R.F. Cain. 1975. Checklist of the
mosses of Ontario. National Museums of Canada, Publi-
cations in Botany 5. 67 pp.
Kershaw, K. A. 1974. Studies on lichen-dominated sys-
tems. X. The sedge meadows of the coastal raised beaches.
Canadian Journal of Botany 52(8): 1947-1972.
Kucyniak, J. 1948. Sur trois additions a la flore bryolo-
gique du Québec. Annales de !1ACFAS 14: 67.
Kucyniak, J. 1949. On the occurrence of Tortula latifolia
Bruch in Québec. Bryologist 52(1): 32-33.
Kucyniak, J. 1950a.. Mise au point sur deux muscinées
québécoises: Hygrohypnum smithii et Oncophorus virens
var. serratus. Annales de TACFAS 16: 154-156.
Kucyniak, J. 1950b. Muscinées nouvelles pour le Québec:
Calypogeia fissa, Odontoschisma elongatum, Didymodon
rufus et Hypnum bambergeri. Naturaliste Canadien
77(1 1-12): 305-312.
Kucyniak, J. 1950c. Le genre Splachnum dans le Québec.
Revue Bryologique et Lichénologique 20(1—2): 38-42.
Kucyniak, J. 1952a. The occurrence of Grimmia teretiner-
vis Limpr. in North America, with notes on its distribu-
tion. Bryologist 55(1): 35-47.
Kucyniak, J. 1952b. Notes sur les Pohlia du Québec. I.
Deux espéces peu fréquentes: P. filiformis et P. proligera.
Naturaliste Canadien 79(6—7): 233-238.
Kucyniak, J. 1954. Notes sur les Pohlia du Québec. Il. P.
bulbifera et P. drummondii. Naturaliste Canadien 81(10-
11): 197-202.
Kucyniak, J. 1955a. Mousses nouvelles ou intéressantes de
la bryoflore du Québec. Annales de ?ACFAS 21: 104-108.
1979
Kucyniak, J. 1955b. Le genre Seligeria dans le Québec.
Annales de PACFAS 21: 110-115.
Kucyniak, J. 1955c. An overlooked moss in the Quebec
flora: Drepanocladus brevifolius. Svensk Botanisk Tids-
skrift 49(1-2): 325-328.
Kucyniak, J. 1955d. Précisions sur le genre Thelia dans le
Québec. Naturaliste Canadien 82(2-3): 45-48.
Kucyniak, J. 1957a. Notes sur les Pohlia du Québec. III.
Addition du P. sphagnicola et extension d’aire du P.
bulbifera. Naturaliste Canadien 84(12): 249-253.
Kucyniak, J. 1957b. Un Fissidens nouveau pour le Québec.
Annales de ! ACFAS 23: 94.
Kucyniak, J. 1958a. Notes sur les Pohlia du Québec. IV.
Ré-intégration du P. cucullata dans notre flore. Natur-
aliste Canadien 85(4): 94-100.
Kucyniak, J. 1958b. Une mousse inattendue pour le Qué-
bec: Prerigoneurum ovatum. Naturaliste Canadien 85(10):
217-224.
Kucyniak, J. 1958c. On Drepanocladus badius in continen-
tal North America. Bryologist 61(2): 124-132.
Kucyniak, J. 1961. Un autre Desmatodon pour le Québec.
Naturaliste Canadien 88(6—7): 161-165.
LeBlanc, F. 1949. Additions a la bryoflore du Québec.
Naturaliste Canadien 76(8-10):; 223-228.
LeBlanc, F. 1951. Ulota drummondii (Hook, et Grev.)
Brid. en Amérique du Nord. Revue Bryologique et Liché-
nologique 20(3—4): 260-262.
LeBlanc, F. 1954. Additions a la bryoflore du Québec (2e
série). Naturaliste Canadien 81(3—4): 90-99.
LeBlanc, F. 1963. Notes sur les mousses du Québec. I.
Naturaliste Canadien 90: 41-50.
NOTES
433
Lepage, E. 1947. Les lichens, les mousses et les hépatiques
du Québec et leur réle dans la formation du sol arable dans
la région du bas de Québec de Lévis 4 Gaspé. Naturaliste
Canadien Extract (fascicle 4. Les mousses): 125-283.
Lewinsky, J. 1974 (1975). The genera Leskeella and Pseu-
doleskeella in Greenland. Bryologist 77(4): 601-611.
Love, D., J. Kucyniak, and G. Johnston. 1958. A plant
collection from interior Quebec. Naturaliste Canadien 85:
25-69.
Macoun, J. and N. C. Kindberg. 1892. Catalogue of Cana-
dian plants. Part 6, Musci. Geological Survey of Canada.
Montreal. 295 pp.
Masson, P. 1967. Notes sur quelques muscinées du Québec.
Annales de PACFAS 33: 39.
Mogensen, G.S. 1973. A revision of the moss genus
Cinclidium Sw. (Mniaceae Mitt.) Lindbergia 2: 49-80.
Steere, W. C. 1947. 6. Musci. Jn Botany of the Canadian
Eastern Arctic. Part II. Thallophyta and Bryophyta.
Edited by N. Polunin. National Museum of Canada
Bulletin 97: 370-490.
Steere, W. C. 1978. The mosses of Arctic Alaska. Bryo-
phytorum Bibliotheca 14. 508 pp.
Vitt, D. H. 1973. A revision of the genus Orthotrichum in
North America, north of Mexico. Bryophytorum Bibli-
otheca |. 208 pp.
Williams, C. 1968. New and additional moss records for
Ontario. Bryologist 71(3): 282-284.
Received 5 March 1979
Accepted 31 March 1979
Swans Wintering on Vancouver Island, 1977-1978
RICHARD W. MCKELVEY
Canadian Wildlife Service, P.O. Box 340, Delta, British Columbia V4K 3Y3
McKelvey, Richard W. 1979. Swans wintering on Vancouver Island, 1977-1978. Canadian Field-Naturalist 93(4): 433-436.
An aerial survey of swans wintering on Vancouver Island, British Columbia in February 1978 revealed a minimum of 1065
birds. Most were believed to be Trumpeter Swans (Olor buccinator) although some Mute Swans (Cygnus olor) were present
on southern Vancouver Island. Cygnets accounted for 21.5% of the total. Changes in wintering distribution were noted: the
largest population shift was into the agricultural areas of Comox and Nanaimo.
Key Words: Trumpeter Swan, Olor buccinator, Mute Swan, Cygnus olor, wintering, Vancouver Island, estuaries.
Swans winter along the coast of western North
America, from Alaska to southern Washington
(Bellrose 1976). Counts of winter populations have
generally been sporadic owing to the extent and
remoteness of the winter habitat, except on Van-
couver Island. Estimates of the number of swans
wintering there have been made periodically over the
last eight years. In 1977-1978 I conducted an aerial
survey of Vancouver Island areas frequented by
swans. Objectives of this survey were further to assess
winter swan populations, determine the proportion of
cygnets, and document changes in major concentra-
tion areas.
Methods
The survey, conducted 16-18 February 1978,
covered most areas previously surveyed (D. Trethe-
wey, personal communication) including most estu-
aries and lakes found to be ice-free. Northern
Vancouver Island, from Cape Scott to Port Hardy,
434
was not surveyed because of poor weather: its
omission is not considered important, because previ-
ous surveys recorded few swans there and habitat is
limited.
A float-equipped Cessna 185 was flown at an
altitude of approximately 215 m. Observations were
made from both sides of the aircraft. In areas of high
concentration the flocks were counted as the aircraft
circled, and the counts were verified later from
photographs.
Results and Discussion
Population Status
Table | compares results of the 1977-1978 survey
with those of previous surveys. Vancouver Island has
been divided into eight regions based on landforms
(after Holland 1964). The regions and sighting
locations are shown in Figure 1.
In the present study 1065 swans were counted, a
result which is not substantially different from those
reported previously. Smith and Blood (1972) esti-
mated a peak population of 1076 in the period
1969-1971. This was based on the results of a single
aerial count of 1013 birds in February 1971, and
several estimates made by others (I. Smith and D.
Trethewey. unpublished data). I. Smith and D.
Trethewey recorded only 892 swans in 1972-1973. The
NUMBER OF SWANS
e 1-10
@ \i\-25
@ 26-50
@ 51-100
KILOMETERS
20 0 20 40 60 80 100
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 1—Swans counted during aerial surveys on Van-
couver Island
Number of swans (cygnets)
Region 1970-71! 1972-73 1977-78
Parksville-Victoria 62 48(4) 2292(45)
Barkley Sound 90 35(1) 109 (30)
Alberni Basin 1128 195(25) 97 (12)
Ucluelet-Quatsino 308 207(27) 166 (27)
Cape Scott 107 90(10) 44 (6)
Kelsey Bay 140 129(20) 76 (38)
Comox Valley 112 108(7) 246 (52)
Adults and immature 760 789 836
cygnets 2463 103 229
'From Smith and Trethewey (unpublished data).
2Including two Mute Swans.
3Based on an average of 24.3% cygnets, after Smith and
Blood (1972).
variability between counts 1s reduced, however, when
cygnets are eliminated from the comparison(Table 1).
There seems to have been little or no increase in
numbers of adult and subadult swans since 1969-
1971. The estimate by Smith and Blood (1972) of a
129% increase over the 1969-1971 period may have
resulted from incomplete surveys, or from the
VANCOUVER
ISLAND
FIGURE |. Locations of swan sightings in the 1977-1978 winter survey.
1979
inclusion of cygnets in their calculations. The year-to-
year variation in cygnet production can be large. so
estimates of population increases may be biased if
cygnet numbers are included.
Much of the apparent increase on this survey may
be the result of previous surveys not having con-
sidered resident Mute Swans (Cygnus olor) estab-
lished locally from escaped park birds. I believe most
swans I observed were Trumpeter Swans (Olor
buccinator) as did Smith and Blood (1972). Both
Mute Swans and Whistling Swans (O. columbianus),
however, are regularly recorded on Vancouver Island
in winter (British Columbia Provincial Museum sight
records (BCPM)).
Records of Mute Swans on Vancouver Island
extend back to at least 1945 (BCPM). The feral
population had increased to at least 100 birds by 1972
(Tatum 1973). Christmas Bird Counts indicate that -
the population is still increasing but is confined to the
Victoria-Ladysmith area (American Birds, Volume
25-32). Although only two Mute Swans were iden-
tified in the 1977-1978 survey, on Somenos Lake near
Duncan, it seems likely that many more of the swans
recorded in that area were of this species.
Few swans were reported in previous aerial surveys
between Ladysmith and Victoria (Table |). Because it
is unlikely that Mute Swans were identified and
excluded from the reported results of those surveys,
they may have been concentrated in areas not
thoroughly surveyed, such as Greater Victoria (D.
Trethewey, personal communication). If Mute Swans
in the Ladysmith-Victoria area were missed in
previous surveys, their inclusion in my survey may be
responsible for the apparent increase. The 65 adults I
counted in that area is close to the number of Mute
Swans reported (67) in the 1976-1977 Christmas Bird
Count (Weseloh 1977, p. 457). Lowering the adult and
subadult total for 1977-1978 by 65 gives a figure very
close to those for 1970-1971 and 1972-1973 (Table 1).
Weekly counts of swans at Comox Harbour and
Port Alberni in 1977-1978 showed 1-3% were Whis-
tling Swans. Other records (BCPM) indicate that Whis-
tling Swans are usually seen only on southern
Vancouver Island. That may be because most ob-
servers are from the southern part of the island, or it
may reflect the more southerly winter distribution of
Whistling Swans (Bellrose 1976). If Whistling Swans
were uniformly distributed a conservative estimate of
their numbers would be 2% or 20 birds.
Cygnets accounted for 229 or 21.5% of the swans
counted in this study. The proportions of cygnets
reported by Smith and Blood (1972) ranged between
22% and 26%, which they suggested was indicative of
an expanding population not yet limited in breeding
habitat. Although productivity between years 1s
NOTES
435
variable (Table 1), factors other than availability of
breeding habitat may be limiting the population. King
(1976) reported a yearly breeding population growth
of only 3.5%. Mortality may be high during the winter
(J. King, personal communication) or during spring
migration in order for the population to be expanding
so slowly.
Major Concentrations
Large concentrations in 1977-1978 were at Nanai-
mo Harbour (58), Mitchell Lake (55), Nanaimo Lakes
(24), Somenos Lake (28), Cowichan River estuary
(35), Port Alberni (72), Sarita River estuary (53).
Kelsey Bay (76), Northy Lake (91), and Comox
Harbour (92). Areas of major use seem to have
changed between surveys (Table 2). The largest
decrease was at Holberg, from 60 in 1970-1971 to only
8 in 1977-1978, but no regular ground counts exist for
that area: local movements may have affected the
counts in one or more surveys.
TABLE 2—Numbers of swans wintering in areas reported to
have large concentrations in previous surveys
Number of swans
Area 1969-71' 1972-732 1977-783
Kelsey Bay 140 129 76
Pt. Alberni 83 195 97
Kyuquot Sound 81 39 35
Tlupana Inlet 78 57 8
Comox Harbour 74 79 185(274)4
Holberg 50 37 8
Herbert Inlet 51 4 1]
Nanaimo 51 48 15]
Chewat River 48 45 28
‘Smith and Blood (1972).
2Smith and Trethewey (unpublished data).
3This study.
4Maximum observed in weekly ground counts at Comox.
1977-1978.
The largest increases have been in the Comox and
Nanaimo areas. Both locations have estuaries, but the
major surrounding land use is agricultural. More
swans were seen on adjacent farmland than on either
estuary in this study. In both areas there has been a
trend in the last 10 yr towards more intensive dairy
farming. Consequently pastures are now planted with
fast-growing grasses of high nutrient content. I believe
the attractiveness of these pastures and their prox-
imity to open water are the main reasons swans are
concentrating in these areas. Although adjacent
estuaries continue to be important when snow or frost
keeps the swans off the pastures, conflicts with
farmers seem inevitable, should swan use of pasture
land continue to increase.
436
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 3—Numbers of swans wintering in areas subject to frequent ground counts or regular Christmas Bird Counts
Location
Date Comox! Nanaimo? Kelsey Bay? Port Algerni3
1972-73 5] 23 No data No data
1973-74 56 14 98 No data
1974-75 144 28 83 No data
1975-76 No data 36 73 178
1976-77 No data 50 59 No data
1977-78 264 75 75 80
'Data from D. Trethewey (personal communication) and this study.
2Data from American Birds. Vol. 27-32.
3Data from Copland (1976. p. 212) for 1974-75. and this study in 1977-78.
Other data support my belief that swans may be
relocating into certain areas (Table 3). Ground counts
conducted by the Canadian Wildlife Service at
Comox showed that peak counts have increased. Data
from Christmas Bird Counts, while not as rigorous as
the ground counts at Comox, show similar trends.
Nanaimo also showed a general increase, while Kelsey
Bay showed a decrease in swan use. Local British
Columbia Fish and Wildlife Branch personnel also
believe that these trends are real: the locations of
major concentrations of swans are changing.
More research is needed to determine the habitat
and food requirements of wintering swans. Alter-
native means of reducing swan use of pasture grasses,
including lure crops and scaring, willalso need study if
such use increases.
Acknowledgments
The assistance of D. Smith, B. Whitehead (pilot),
D. Trethewey, R. W. Campbell, J. P. Kelsall, and the
reviewers is greatly appreciated.
Literature Cited
Bellrose, F. C. 1976. The ducks, geese and swans of North
America. Stackpole Books, Harrisburg, Pennsylvania.
Copland, H.W.R. (Compiler). 1976. Christmas bird
count for 1975-76; Western Canada. American Birds 30:
208-216.
Holland, S.S. 1964. Landforms of British Columbia. A
physiographic outline. British Columbia Department of
Mines and Petroleum Resources, Bulletin 48.
King, J. G. 1976. The current status and future of the
Alaska Trumpeter Swan population. Proceedings of the
Fifth Trumpeter Swan Society Meeting (in press).
Smith, I. D. and D. A. Blood. 1972. Native swans winter-
ing on Vancouver Island over the period 1969-71. Can-
adian Field-Naturalist 86: 213-216.
Tatum, J.B. 1973. Annual bird report for southern Van-
couver Island 1972. Victoria Natural History Society.
Victoria, British Columbia.
Weseloh, D. V.(Compiler). 1977. Christmas bird count for
1976-77: Western Canada. American Birds 31: 457-466.
Received 18 September 1978
Accepted 27 March 1979
Interspecific Vocal Mimicry by Pine Grosbeaks
PETER TAYLOR
P.O. Box 597, Pinawa, Manitoba ROE 1L0
Taylor, Peter. 1979. Interspecific vocal mimicry by Pine Grosbeaks. Canadian Field-Naturalist 93(4): 436-437.
Mimicry of the calls of the Hairy Woodpecker, Gray Jay, American Robin, and Redpoll (sp.) by adult male Pine Grosbeaks,
during “whisper singing” on the species’ winter range, is described.
Key Words: Pine Grosbeak; Pinicola enucleator; vocal mimicry.
This note describes vocal mimicry during “whisper
singing” by adult male Pine Grosbeaks (Pinicola
enucleator) on the species’ winter range in south-
eastern Manitoba. Interspecific vocal mimicry has
been reported for several bird species (e.g., Adkisson
and Conner 1978; Armstrong 1973), but the only
suggestion of such behavior by Pine Grosbeaks is a
reference to four birds uttering “an amplified version
of the common flight call of the goldfinch” (Bent 1968,
p. 333). Soft, warbled “whisper singing” is docu-
1979
mented for many North American fringillids in-
cluding Pine Grosbeaks (Bent 1968).
At 09:00 on 26 February 1978 I noted several Pine
Grosbeaks in Pinawa; three of these were uttering soft
warbling songs. One adult male sang particularly
softly, and interspersed the warbled phrases with
realistic imitations of several other birds’ calls, uttered
clearly but softly. | approached within 5 m of the bird
as it perched alone in a small aspen, so there is no
doubt that it gave all the calls noted. These included
the ‘pic’ call-note of a Hairy Woodpecker ( Picoides
villosus), the harsh *kuk-kuk-chikkikik’ call of a Gray
Jay (Perisoreus canadensis), the clucking call and
rattling alarm-cry of an American Robin (Turdus
migratorius), and the call-note, with rising inflection,
of a Redpoll (Carduelis sp.). Some slurred whistles
were Starling-like (Sturnus vulgaris), but not un-
equivocally recognizable. A muted version of the Pine
Grosbeak’s own triple-noted flight call was also
included.
These various calls were interjected, with no
apparent ordered sequence, at intervals of about 3 s
into the warbled song, and the medley was sustained
for periods of 20 s or more. All of the imitations were
not included in each bout of singing. The bird sang for
about half of the observation period of about 15 min,
and was still singing when I left.
At 11:30 on 5 March 1978 I heard what was
presumably the same bird uttering similar vocaliza-
tions at the same location. The song was less sustained
on this occasion, and interference from extraneous
noise made observation difficult.
NOTES
437
At 12:00 on 4 February 1979 I observed an adult
male Pine Grosbeak uttering a “whisper song” while
foraging in a spruce tree in Pinawa. This song
continued, with one 15-s break, for about 6 min. It
consisted of a variety of whistled and warbled phrases,
and included imitations of American Robin and
Hairy Woodpecker call-notes.
Although it is possible that the same individual Pine
Grosbeak was involved in both the 1978 and 1979
records, the observation of this behavior in two
winters does suggest that vocal mimicry may not be
uncommon in Pine Grosbeaks. All of the birds
imitated, except the American Robin, winter regu-
larly in the Pinawa area. If the Pine Grosbeaks had
wintered in southeast Manitoba, then they would
have learned the robin calls the previous fall at the
latest.
I thank Richard W. Knapton for helpful comments
on this note.
Literature Cited
Adkisson, C.S. and R.N. Conner. 1978. Interspecific
vocal imitation in White-eyed Vireos. Auk 95: 602-606.
Armstrong, E. A. 1973. A study of bird song. Dover, New
York, New York. 347 pp.
Bent, A.C. (and collaborators). 1968. Life histories of
North American cardinals, grosbeaks, buntings, towhees,
finches, sparrows and allies. U.S. National Museum
Bulletin 237.
Received 14 November 1978
Accepted 3 February 1979
7 May 1979 (amended)
Influence of Weather on Aggression in Tree Swallows!
REID N. HARRIS
Department of Zoology, University of Maryland, College Park, Maryland 20742 USA
Harris, Reid N. 1979. Influence of weather on aggression in Tree Swallows. Canadian Field-Naturalist 93(4): 437-438.
Cool foggy weather significantly reduced territorial aggression in a Tree Swallow (/ridoprocne bicolor) population in New
Brunswick.
Key Words: aggression, weather, Tree Swallow, /ridoprocne bicolor.
Weather can influence the behavior of animals in
nature. For example, Grubb (1978) and Travis (1977)
have found that weather affected the daily and
seasonal foraging habits of birds. This note reports on
the effects of weather on territorial aggression in the
'This paper is a contribution from the Bowdoin Scientific
Station, Kent Island, New Brunswick, Canada.
Tree Swallow, /ridoprocne bicolor.
Study Site and Methods
From 17 May until | August 1978, I conducted a
study of aggression, territoriality, and reproductive
success in Tree Swallows on Kent Island, New
Brunswick, Canada. Tree Swallows have nested in
uniformly spaced man-made boxes in two open fields
on Kent Island for 45 yr (Paynter 1954). In 1978, two
438
new spatial patterns were created by moving nest-
boxes from their original 30-m uniform spacing,
leaving 12 uniformly spaced boxes as a control
comparable with previous years. The first spatial
pattern consisted of alternating, at 30-m intervals, a
pair of boxes, | m apart, with a single box. Twenty-
eight boxes were arranged in this manner. The second
pattern consisted of two replicates of a spiral
arrangement of five boxes placed so that boxes were
1 m,2 m,4 m,and 8 maway from the central box. The
two spirals were 30 m apart: also 30 m separated the
nearest boxes of adjacent spatial patterns.
All breeding females and several breeding males
were color-marked for individual recognition (see
Hoogland and Sherman 1976). Adult Tree Swallow
behavior was recorded for 4 wk, including the
approximately 20-d nestling period of Tree Swallows
on Kent Island (Paynter 1954: this study). Twenty-five
occupied nest-boxes from all spatial patterns were
selected at random (by drawing numbers from a hat)
from a total of 36 active boxes for observation. Two or
three active nest-boxes were watched simultaneously
for 45-min periods. Each box was observed for about
7h during the 4-wk observational study: a total of
182 h of observation was accumulated. Observations
were conducted in all kinds of weather except driving
rain from 09:00 to 14:30 Atlantic Daylight Time.
Warm days (>12°C) with high visibility (>100 m)
were classified as “weather 1” days. Cool days
(S<12°C) with low visibility (>100 m:; i.e., fog) were
Classified as “weather 2” days. All observational data
fit into one of these categories: foggy days were always
cool.
During observations, the amount of territorial
aggression and the locations of territorial boundaries
were carefully monitored. Fights were defined as two
or more birds actually contacting one another in an
aggressive territorial encounter. Chases were similar
to fights, but lacked actual contact. All aggression
occurred in the context of territorial defense of nest-
boxes, 1.e., marked and unmarked resident birds of
both sexes excluded unmarked swallows from their
territories throughout the nestling period.
Results and Discussion
Cool foggy weather significantly reduced territorial
aggression in Tree Swallows (Table 1). One striking
example of the influence of weather on aggression was
observed on 11 July. From the beginning of observa-
tions at 09:00 until 10:00, cold foggy weather
prevailed, and aggression was non-existent. At 10:00,
a rapid clearing began. Concomitantly, swallows at
the two active boxes being watched began defending
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE I—Aggression rates, mean + SE of aggressive inter-
actions per pair of Tree Swallows during a 45-min period, by
weather are based on 182 box h. All 25 watched pairs were
observed on weather | days, but only 19 pairs on weather
2 days
Aggression Weather | Weather 2 ies
Fights AA) 22 O22 0.18+0.16 <0.001
Chases 0.61 + 0.28 0.11 + 0.03 >0.05
Total 2.6 + 0.43 0.29+0.07 <0.001
‘t-test; variances unequal.
their nest-boxes against unmarked intruders. Fifteen
separate fights that involved actual contact between
birds were observed between 10:10 and 10:30. Aggres-
sive activity was directly associated with weather
conditions.
The exact proximal causes of the reduction in
aggression during cool foggy weather are unknown.
Lowered visibility associated with fog, however, could
interfere with the swallows’ cues of individual recogni-
tion so that potential aggressors may not be recog-
nized except at very close distances. Alternatively,
during periods of low visibility, foraging for the same
quantity of insects must have taken longer than
foraging in clear weather. Under foggy conditions,
and given a parental commitment to feed nestlings,
most of the swallows’ time would have been taken
foraging. Whether these explanations can be distin-
guished or demonstrated is the subject for future
work.
Acknowledgments
This work was supported in part by the A. O. Gross
Fund. I thank D. E. Gill, G. L. Holroyd, P. Wood-
ward, and an anonymous referee for helpful com-
ments on the manuscript, and R. Robertson for
suggesting the spiral pattern for spacing nest-boxes.
Literature Cited
Grubb, T. C. 1978. Weather-dependent foraging rates of
wintering woodland birds. Auk 95: 370-376.
Hoogland, J.L. and P. W. Sherman. 1976. Advantages
and disadvantages of Bank Swallow coloniality. Ecologi-
cal Monographs 46: 33-58.
Paynter, R.A. 1954. Interrelations between clutch size,
brood size, prefledging survival and weight in Kent Island
Tree Swallows. Bird-Banding 25: 35-58, 102-111, 136-149.
Travis, J. 1977. Seasonal foragingina Downy Woodpecker
population. Condor 79: 371-375.
Received 19 February 1979
Accepted 11 April 1979
1979 NOTES 439
Abandoned Windmill Used as a Nesting Site
by Great Blue Herons
JEAN-LUC DESGRANGES
Canadian Wildlife Service, 2700 Laurier Blvd., P.O. Box 10100, Ste-Foy, Québec G1V 4H5
DesGranges, Jean-Luc. 1979. Abandoned windmill used as a nesting site by Great Blue Herons. Canadian Field-Naturalist
93(4): 439-440.
An active Great Blue Heron (Ardea herodias) nest was discovered on the platform of an abandoned windmill at Batture aux
Loups-Marins, Quebec. Among the hundred or so heronries known in Quebec, this was the only one where a man-made
structure was used.
Un nid actif de Grand Héron (Ardea herodias) installé sur la plateforme d’une éolienne désaffectée a été découvert a la Batture
aux Loups-Marins, Québec. Parmi la centaine de héronniéres connues au Québec jusqu’a ce jour, cest la premiére fois que
Yon observe utilisation d’une structure artificielle par cette espéce.
Key Words: Ardea herodias, nesting platform.
On 25 April 1978, while conducting anaerialsurvey —_ any of the nests located in trees, the usual nesting site
of aquatic bird colonies in the St. Lawrence estuary for this species.
for the Canadian Wildlife Service, Pierre Dupuis and
I discovered an active Great Blue Heron (Ardea
herodias) nest on the platform of an abandoned
windmill at Batture aux Loups-Marins (47°14’N,
70°25’W) (Figure 1). We located four other nests
being built in a grove of tall willows (Sa/ix sp.) in the
vicinity of the windmill. There were no Great Blue
Herons at this site prior to 1978 (Reed 1973; personal
observation 1977).
On | June, the heronry contained eight active nests,
four of which contained only eggs (K + SD = 3.0 + 1.6)
whereas the others, including the nest on the windmill,
held both eggs and young. On 3 July, six nests
remained, all with young herons (K + SD = 2.0 + 1.2).
The nest-located on the windmill held four young
herons whereas the others held fewer than three each.
Although Great Blue Herons have been recorded
nesting on duck-hunting blinds (Stotts 1959) and
navigational buoys (Henny 1978), it is nonetheless
unusual for them to nest on artificial platforms.
Among the hundred or so heronries known in Quebec,
this was the only one in which a man-made structure
was used (DesGranges, unpublished data). Neverthe-
less, other species of herons have been known to nest on
artificial structures (MclIlhenny 1934; Finkenstaedt
and Heckenroth 1974; Wiese 1976). This suggests that
the erection of elevated platforms where Great Blue
Heron natural nest sites have been destroyed could
prove to be an effective remedial technique, providing
other features of the habitat remain suitable. At
Batture aux Loups-Marins, the artificial platform was met
occupied from the time the heronry was first establish- Figure 1. Great Blue Heron nest built on platform of
ed and the nest found on it produced more young than abandoned windmill.
I thank Transport Canada for making a helicopter
available for our use in the St. Lawrence estuary
survey.
440
Literature Cited
Finkenstaedt, C. and H. Heckenroth. 1974. Eine kunstliche
kolonie grundung beim Graureiher (Ardea cinerea).
Vogelwelt 95: 227-230.
Henny, C. D. 1978. Great Blue Herons respond to nesting
habitat loss. Wildlife Society Bulletin 6: 35-37.
Mclihenny, E. A. 1934. Bird city. Christopher Publishing
House, Boston. 203 pp.
Reed, A. 1973. Aquatic bird colonies in the St. Lawrence
THE CANADIAN FIELD-NATURALIST
Vol. 93
estuary. Service de la faune du Québec, Bulletin 18. 54 pp.
Stotts, V. D. 1959. Offshore duck blinds: Their use by wild-
life and how to improve them for wildlife use. Maryland
Conservationist 36: 23-26.
Wiese, J.J. 1976. Courtship and pair formation in the
Great Egret. Auk 93: 709-724.
Received 8 January 1979
Accepted 9 June 1979
Unusually Late Pregnancy of a Muskrat in Southeastern
New Brunswick
G. R. PARKER
Canadian Wildlife Service, P.O. Box 1590, Sackville, New Brunswick E0OA 3C0
Parker, G. R. 1979. Unusually late pregnancy of a Muskrat in southeastern New Brunswick. Canadian Field-Naturalist
93(4): 440-441.
A Muskrat (Ondatra zibethicus zibethicus) containing nine embryos was trapped on 6 November 1978 on the Tantramar
marshes in southeastern New Brunswick. The projected birth date for the embryos was the last week of November. This is the
latest reported pregnancy for a muskrat in eastern Canada.
Key Words: Muskrat, late pregnancy, New Brunswick, Ondatra zibethicus zibethicus.
As part of a study of Muskrats (Ondatra zibethicus
zibethicus) on the Tintamarre National Wildlife Area,
Westmorland County, New Brunswick, carcasses of
Muskrats caught during the 1978 fall trapping season
(4 November — 31 December) were examined for sex,
age, productivity, and other morphological measure-
ments. Productivity was measured by placental scar
counts from macroscopically examined female uterl.
The uterus of one female caught on 6 November
contained nine embryos; these averaged 5-7 mm in
length and were believed to be approximately 10-14 d
old. The normal gestation period for Muskrats in
northeastern North America is 25-30 d (Errington
1963). This Muskrat would have given birth during
the last week of November.
This Muskrat had been live-trapped and ear-tagged
2 August; it weighed 1000 g. When caught in
November it weighed 1490 g. Its exact age is
uncertain. When tagged in August, it was subjectively
classified as an adult, based upon size and appearance.
When examined in November the molar fluting was at
bone line, which left the age of the animal in question.
The uterus showed no other scars of earlier preg-
nancies.
It seems most reasonable that this Muskrat was
born late in the autumn of 1977, and that possibly it
too came from a fall litter. This would account for its
adult appearance in August, the questionable age
from molar fluting, the absence of previous breeding,
and its substantial weight gain in late summer and fall.
The Tantramar marshes of the New Bruns-
wick — Nova Scotia border region con-
tain some of the best habitat for Muskrats in the
Maritimes. Under favorable conditions, it seems
reasonable that autumn or fall breeding may oc-
casionally occur, and that some kits survive the winter
to enter the next year’s breeding population. Such
late-born females, however, probably do not breed
until late the following summer.
Muskrats may give birth during all months of the
year in California (Dixon 1922), Louisiana (Svihla
and Svihla 1931), and Texas (Lay 1945). Further
north, in Maryland, breeding normally ceases by late
October (Forbes 1942). Late litters have been reported
in September from Wisconsin (Beer 1950), east
Tennessee (Schacher and Pelton 1975), Connecticut
(Smith and Jordan 1976), and Manitoba (McLeod
and Bondar 1952), and in October from Idaho
(Errington 1963). In New Brunswick (Dilworth 1967)
and Prince Edward Island (Dibblee 1970) the latest
observed litters were born in September. Errington
suggested that it is quite possible litters might rarely be
sired during winter in lowa. This specimen from the
Tantramar marshes of New Brunswick, however, is
the first documented evidence of a November preg-
nancy of a Muskrat in eastern Canada.
1979
Literature Cited
Beer, J. R. 1950. The reproductive cycle of the Muskrat
in Wisconsin. Journal of Wildlife Management 14(2):
151-156.
Dibblee, R. L. 1970. The reproduction and productivity of
Muskrats on Prince Edward Island. Proceedings of the
Canadian Society of Wildlife and Fisheries Biologists,
Atlantic Chapter, Fredericton, November 4-6. pp. 114—
122.
Dilworth, T. G. 1967. The life history and ecology of the
Muskrat under severe water level fluctuations. Proceed-
ings of the Canadian Society of Wildlife and Fisheries
Biologists, Atlantic Chapter, Charlottetown, November
28-29. 17 pp.
Dixon, J. 1922. Rodents and reclamation in the Imperial
Valley. Journal of Mammalogy 3(3): 136-146.
Errington, P. L. 1963. Muskrat populations. Iowa State
University Press, Ames. 665 pp.
Forbes, T.R. 1942. The period of gonadal activity in the
Maryland Muskrat. Science 95: 382-383.
NOTES
44]
Lay, D. W. 1945. Muskrat investigations in Texas. Journal
of Wildlife Management 6(1): 56-76.
McLeod, J. A. and G. F. Bondar. 1952. Studies on the
biology of the Muskrat in Manitoba. Part I. Oestrous
cycle and breeding season. Canadian Journal of Zoology
30(4): 243-253.
Schacher, W. H. and M.R. Pelton. 1975. Productivity in
Muskrats in East Tennessee. Proceedings of the 29th
Southeast Game and Fish Commission, St. Louis, Mis-
souri, October 12-15. 26 pp.
Smith, H.R. and P.A. Jordan. 1976. An unexploited
population of Muskrats with unusual biomass, produc-
tivity, and body size. State Geological and Natural
History Survey of Connecticut, Report of Investigations
Number 7. 16 pp.
Svihla, A. and R. D. Svihla. 1931. The Louisiana Muskrat.
Journal of Mammalogy 12(1): 12-28.
Received 11 January 1979
Accepted 2 May 1979
Lynx Movements and Habitat Use in Montana!?
GARY M. KOEHLER, MAURICE G. HORNOCKER, and HOWARD S. HASH
Idaho Cooperative Wildlife Research Unit, University of Idaho, Moscow, Idaho 83843
Koehler, Gary M., Maurice G. Hornocker, and Howard S. Hash. 1979. Lynx movements and habitat use in Montana.
Canadian Field-Naturalist 93(4): 441-442.
Movements of two Lynx (Lynx canadensis) were monitored by radio telemetry: an adult male for 74 mo from March to
October and an adult female during January. Home range area for the male, determined from 21 radio locations, was 36 km2.
Most locations were in densely stocked stands of Lodgepole Pine (Pinus contorta) resulting from the 1910 fires. Snowshoe
Hares (Lepus americanus), their principle prey, were also most abundant in these stands.
Key Words: habitat, Lynx (Linx canadensis), Snowshoe Hare (Lepus americanus), activity patterns.
Little is known about seasonal habitat use or home
range size of Lynx (Lynx canadensis). Most previous
studies used snow-tracking techniques (Saunders
19632) Nelhis et al. 1972; Brand) et al. 11976): Berrie
(1973) used radio telemetry to determine home range
size in Alaska. In conjunction with a Wolverine (Gu/o
gulo) study in northwest Montana, we monitored
movements of two Lynx seasonally during 1977 by
radio telemetry.
Methods
Twenty live-traps were set from January through
‘Contribution of the Idaho Cooperative Wildlife Research
Unit; the United States Fish and Wildlife Service, the Idaho
Department of Fish and Game, the University of Idaho, and
the Wildlife Management Institute cooperating.
*University of Idaho College of Forestry, Wildlife and Range
Sciences Publication No. 168.
April along 40 km of Forest Service trail. Traps,
measuring 40 X 40 X 65 cm, were constructed of tubu-
lar steel frames, chain link fence sides, and sliding steel _
doors. Lynx were immobilized with ketamine hydro-
chloride (“Ketalar,” Parke Davis) in dosages approxi-
mating 21 mg/kg body weight. Lynx were eartagged
and tattooed in the lip and on the body under the
foreleg, measured, evaluated for general physical
condition, and fitted with collars containing radio
transmitters. Radio signals were monitored from
fixed-wing aircraft and the ground.
The relative abundance of Snowshoe Hares ( Lepus
americanus) in various forest types was obtained by
counting the number of tracks crossed per kilometre
of trail after fresh snowfall. To help differentiate
tracks only those traveling to the west of the trail and
spaced 3-m apart were recorded. A vegetative descrip-
tion of each section of trapline included dominant
overstory species, age class, and relative density.
442
THE CANADIAN FIELD-NATURALIST
Vol. 93
TABLE 1—Relative abundance of Snowshoe Hare in four vegetative cover types along trapline during 14 d of survey
, Tracks
Distance
Stand age, Stand surveyed Total ty
Vegetative cover yr density (km) no. No. km -d
Lodgepole Pine, pure stands < 80 Dense 9.0 Sil 7/ 2.54
Grassland islands in dense Lodgepole Pine < 80 Dense 37) 96 Daisy
Subalpine Fir-Englemann Mature Medium 2.4 15 0.45
Spruce > 100
Islands of Lodgepole Pine, Douglas Fir,
Ponderosa Pine (Pinus ponderosa) in Mature Sparse 5.6 11 0.14
> 100
grasslands
Results and Discussion
One adult male (weight 10 kg), one adult female
(7 kg), and one juvenile female (4 kg) were captured.
The juvenile accompanied the adult female and was
not radio-collared because it was too small. The adult
female was captured on 14 January and located 8
times prior to being found dead on 31 January. Death
was believed due to predation by a Mountain Lion
(Felis concolor). The male was captured on 10 March
and located 21 times by 27 October.
Most locations for the male and female were in
young densely stocked stands of Lodgepole Pine
(Pinus contorta). Twenty-six of the 29 locations (90%)
occurred in timbered areas burned in 1910 and the
remainder occurred in mature Douglas Fir ( Pseudo-
tsuga menziesii) — Western Larch (Larix occidentalis)
stringers along stream bottom within the 1910 burn.
Of the locations burned in 1910, 23 (88%) occurred on
xeric sites where Lodgepole Pine was dominant and 3
(t2%) on mesic sites where Subalpine Fir (Abies
lasiocarpa) and Englemann Spruce ( Picea engleman-
nii) were dominant. No locations occurred in open
grassland or semi-open areas, 90% were in densely
stocked stands, and 10% in medium-stocked stands.
The estimated home range size for the male was
36 km?. This figure is similar to that found in
other studies. Winter tracking in Newfoundland
showed a home range size between 15.5 and
20.7 km? (Saunders 1963). In Alberta they were from
11.1 to 49.5 km? (Brand et al. 1976). In Alaska, Berrie
(1973) found Lynx to range from 12.8 to 25.5 km2.
Snowshoe Hares were most abundant in densely
stocked stands of Lodgepole Pine (Table 1). Brand et
al. (1976) and Adams (1959) found that hares were
most abundant in dense stands. Winter ground
tracking during this study and in Alberta (Brand et al.
1976) indicate that Lynx concentrate hunting activity
within areas of high hare activity because hares are
their main food (Brand et al. 1976) and 90% of Lynx
radio locations were in these stands. Saunders (1963)
found that Lynx activity and the location of the home
range boundary coincided almost exactly with a tract
of 10- to 20-yr-old growth timber.
Our data suggest that Lynx concentrate activity in
areas of high Snowshoe Hare activity, particularly in
young dense stands of Lodgepole Pine. Radio tele-
metry indicates that there is no change in range areas
or habitat use throughout the seasons.
Acknowledgments
The United States Forest Service provided field
facilities. The cooperation of D. Minister, D. Owen,
and K. Granrud of the United States Forest Service is
sincerely appreciated. We express special thanks to
T. W. Koehler for assisting with the field work.
Financial support was provided by the National
Science Foundation, United States Forest Service,
National Geographic Society, New York Zoological
Society, National Wildlife Federation, Audubon
Society, National Rifle Association, Wildlife Man-
agement Institute, Montana Department of Fish and
Game, and the Boone and Crockett Club.
Literature Cited
Adams, L. 1959. An analysis of a population of Snowshoe
Hares in northwestern Montana. Ecological Monographs
29(2): 141-170.
Berrie, P. M. 1973. Ecology and status of the Lynx in
interior Alaska. Jn The world’s cats. Edited by R.L.
Eaton. Volume |. Proceedings of the International Sym-
posium on the World’s Cats, March 1971. Published by
World Wildlife Safari, Winston, Oregon. pp. 4-41.
Brand, C. J., L. B. Keith, and C. A. Fischer. 1976. Lynx
response to changing Snowshoe Hare densities in central
Alberta. Journal of Wildlife Management 40(3): 416-
428.
Nellis, C. H., S. P. Wetmore, and L. B. Keith. 1972. Lynx-—
prey interactions in central Alberta. Journal of Wildlife
Management 36(2): 320-329.
Saunders, J. J., Jr. 1963. Movements and activities of the
Lynx in Newfoundland. Journal of Wildlife Management
27(3): 390-400.
Received 28 February 1979
Accepted 30 May 1979
1979
NOTES
Decline of Summering Bald Eagles in Central New Brunswick
R. F. STOCEK
The Maritime Forest Ranger School, Fredericton, New Brunswick E3B 4X6
StocekeuRe F:
443-445.
1979. Decline of summering Bald Eagles in central New Brunswick. Canadian Field-Naturalist 93(4):
The numbers of Bald Eagles ( Haliaeetus leucocephalus) that summer in central New Brunswick have decreased considerably.
This reflects chiefly a decline in the proportion of immature birds.
Key Words: Bald Eagle, Haliaeetus leucocephalus, New Brunswick, abundance, age composition.
The Bald Eagle ( Haliaeetus leucocephalus) in New
Brunswick is listed as endangered under the provincial
Endangered Species Act of 1976. It was considered an
uncommon resident and transient by Squires (1976),
who reported that numbers of the bird had decreased
drastically; however, there do not appear to be any
published accounts giving details of that decline. This
paper describes the decrease in numbers and change in
age composition of summering Bald Eagles in the
lower St. John River basin and other parts of central
New Brunswick (between 45°30’N and 47°00’N). The
104-km?2 French Lake district, west of Grand Lake,
Queens County, and centered at 45°50’N, 66°17’W, is
of particular interest here (Figure 1). It includes the
shores of French, Indian, and Maquapit Lakes, the
thoroughfare joining them, and Loders and Porto-
bello Creeks in Sunbury County.
Although distributed over the entire province in the
past, the resident Bald Eagle, H. /. alascanus, was
never considered numerous (Moore 1928). More
recently, banding returns suggested that most sum-
mering eagles belonged to the southern subspecies,
H. |. leucocephalus (Squires 1952). Those birds,
mostly immatures coming from Florida (Broley 1947),
wandered into the Maritime region in April and May,
and returned southward in August and September
(Wright 1953). The wintering birds of the resident
form apparently move inland from coastal areas in
February and March (R. F. Stocek and P.A. Pearce,
unpublished data), and return to the coast late in the
fall (R. F. Stocek, unpublished data).
Methods
Bald Eagle records on file in the Northeastern
Wildlife Station at the University of New Brunswick
and at the New Brunswick Museum were compiled,
with observations from the various naturalist clubs
and provincial and federal resource agencies. Rela-
tively few quantitative data were available prior to
1959 except for the French Lake district records of the
Northeastern Wildlife Station, extending from 1949
to 1962. The French Lake sight records are treated
FiGuRE |. Map of New Brunswick showing the major areas
mentioned in the text. The French Lake district,
shown in dark shading, lies next to Grand Lake in
Queens County. The lower St. John River extends
downstream from Fredericton (the dark circle) and is
shown enclosed by the solid outlines of the four basin
counties. The central part of the province is defined
by the marginal lines of latitude, 45°30’N and
47°00'N.
separately, rather than being combined with the
scattered data from the rest of the province, because
so much more effort was spent on this relatively small
area. Numerous low-level aerial flights (both fixed-
wing and helicopter) were made over the lower St.
John River basin during the spring and summer of
1974 and 1975. As used here, winter refers to
December, January, and February; spring to March,
April, and May; summer to June, July, and August,
and fall to September, October, and November.
444
Results and Discussion
Cumulative sight records between 1959 and 1975
show that central New Brunswick accounted for 55%
of the total spring observations and 60% of the
summer observations of Bald Eagles in the province.
Typically most of the eagles in this region were seen in
the summer (38% of the area total) and spring (30%),
and fewer in the winter and fall (22% and 10%,
respectively).
Thirty years ago the basin of the lower St. John
River was thought to be one of the most important
summer habitats of H. 1. lewcocephalus in the north-
east maritime region of the continent. The peak
summer population of Bald Eagles in the French Lake
district was estimated at 54 in 1949 and 45 in 1950
(Wright 1953). The Foshay Lake - Grimross Neck
area 20 km to the southeast appeared to support a
similar number in 1949. During those years a United
States Fish and Wildlife Service aerial survey team
saw more eagles in this area than in all the rest of the
maritime provinces. Practically all the eagles counted
in New Brunswick were seen here, with as many as 18
in the air at one time near French Lake. A total
population of at least 100 eagles summering in the
lower St. John River basin was suggested (B.S.
Wright, unpublished data). Large numbers of sum-
mering eagles had also been reported there in May and
June, 1937 to 1944, especially along Portobello Creek
and from Fredericton to Jemseg (H. S. Peters, un-
published data). During the 1930s and 1940s, it was
not uncommon to see 20-30 eagles in a day on the
waterfowl breeding grounds in central New Bruns-
wick (Squires 1976).
The mean numbers of summering eagles seen in the
French Lake district declined during the 1950s and
early 1960s. According to data from the North-
eastern Wildlife Station, 2.3 birds were seen per
successful day* during June-July, 1953-1954. By
1961-1962 this figure was down to 1.3; eagles then
were being seen only infrequently by station per-
sonnel. B.S. Wright (personal communication), who
had studied waterfowl in the St. John swamps and
marshes since 1945, believed that there were no more
than 10 eagles on the entire area in the summer of
1964. On | August 1974, I surveyed by helicopter
125 km of shoreline in the French Lake district and
another 179 km of adjacent Grand Lake and Jemseg
River; no Bald Eagles were seen. A_ provincial
naturalist conducting a group canoe trip through the
district and the nearby Oromocto River drainage
reported that only one eagle was seen during the
192-km trip in mid-July 1974. The only eagles I saw
*A day on which at least one eagle was sighted; unfor-
tunately statistics on total effort (days) are not available.
THE CANADIAN FIELD-NATURALIST
Vol. 93
during the many flights I made through the French
Lake area in 1974 and 1975 were the one pair nesting
there.
The decrease of Bald Eagles in certain parts of
North America has been accompanied by a decline in
the proportion of immature birds in those populations
(Sprunt 1969). The percentage of immatures in the
French Lake district summering population for 1953
to 1962 is shown in Table |. The influx of wandering
birds probably accounted for the high May values,
while birds leaving the area in August reduced the
number seen. B.S. Wright (unpublished data), re-
ported a high 1:4 adult to immature ratio for
the district during the spring-summer period in 1949.
TABLE 1—The monthly distribution of immature Bald
Eagles in the French Lake district of New Brunswick from
1953 to 1962, expressed as a percent of the total known-age
birds seen
Month Q% Immature Total no. seen
May 44 4]
June 23} 73
July 2) 70
August 10 19
By 1953-1962 the ratio had changed to 4:1. Values of
30% immatures recorded in 1953-1954 and 13% in
1961-1962 suggested a decline during that period.
Although data are fragmentary for the intervening
years, June and July sightings provide some insight.
From 1953 to 1956, 25% of the 79 eagles seen in the
district were immatures. Station personnel reported
19% immature birds (of 64 seen) from 1959 to 1962.
New Brunswick sight records show that, in the
province as a whole, but excluding data from the
French Lake area, the proportion of immature eagles
seen In spring and. summer decreased from an average
of 32% during 1967-1973 to 22% in 1974-1975
(Table 2). These comparative seasonal values very
likely reflect a real difference between the two time
periods, because the increased effort in 1974-1975
TABLE 2— The seasonal distribution of immature Bald
Eagles in New Brunswick from 1967 to 1973 and
1974-1975, expressed as a percent of the total known-age
birds seen (in parentheses). French Lake district statistics are
not included here
% Immature, % Immature,
Season 1967-1973 1974-1975
Winter 20 (56) 20 (35)
Spring 25 (76) 14 (50)
Summer 37 (104) 27 (93)
Fall 22 (49) 20 (25)
1979
would have been expected to reveal a greater rather
than a smaller proportion of the less conspicuous
immature birds.
The decline in the summering population of Bald
Eagles in central New Brunswick and probably
throughout the province was to be expected. The
decrease in Bald Eagle reproduction in other regions
of North America during the last two decades is also
well documented (e.g., Sprunt 1969; Sprunt et al.
1973). Both Broley (1950, 1958) and Howell (1958)
reported increasing nest failures in Florida, which was
an important source of birds summering in the
Maritimes.
Acknowledgments
I am grateful to the late Bruce Wright, to Peter
Pearce, and John Baird for their helpful suggestions
and discussions. The interest of many individuals
and agencies, and particularly the New Brunswick
Department of Natural Resources in contributing
valuable data, is greatly appreciated. The Canadian
Wildlife Service provided financial support for this
work.
Literature Cited
Broley, C. L. 1947. Migration and nesting of Florida Bald
Eagles. Wilson Bulletin 59: 3-20.
NOTES
445
Broley, C.L. 1950. The plight of the Florida Bald Eagle.
Audubon Magazine 52: 42-49.
Broley, C. L. 1958 The plight of the American Bald Eagle.
Audubon Magazine 60: 162-163, 171.
Howell, J.C. 1958. Further history on some Bald Eagle
nest sites in east-central Florida. Auk 75: 96-98.
Moore, W.H. 1928. A list of the birds of New Brunswick,
Canada. 67th Annual Report, Department of Lands and
Mines 1927: 91-112.
Sprunt, A., IV. 1969. Population trends of the Bald Eagle
in North America. /n Peregrine Falcon populations: their
biology and decline. Edited by Joseph J. Hickey.
University of Wisconsin Press, Madison. pp. 347-351.
Sprunt, A., 1V, W. B. Robertson, Jr., S. Postupalsky, R. J.
Hensel, C. E. Knoder, and F. J. Ligas. 1973. Compara-
tive productivity of six Bald Eagle populations. Trans-
actions of the North American Wildlife and Natural
Resources Conference 38: 96-105.
Squires, W. A. 1952. The birds of New Brunswick. New
Brunswick Museum Monographic Series Number 4.
164 pp.
Squires, W. A. 1976. The birds of New Brunswick. New
Brunswick Museum Monographic Series Number 7.
210 pp.
Wright, B.S. 1953. The relation of Bald Eagles to breeding
ducks in New Brunswick. Journal of Wildlife Manage-
ment li 55—02.
Received 30 October 1978
Accepted 10 May 1979
Lesser Black-backed Gull, Larus fuscus, in Labrador Waters
KEVIN D. POWERS
Manomet Bird Observatory, Manomet, Massachusetts 02345 USA
Powers, Kevin D. 1979. Lesser Black-backed Gull, Larus fuscus, in Labrador waters. Canadian Field-Naturalist 93(4):
445-446.
An adult Lesser Black-backed Gull, Larus fuscus, sighted 140 km E of Nain, Labrador, adds to evidence that the species
occurs in small numbers through eastern North American waters.
Key Words: Lesser Black-backed Gull, Larus fuscus, Labrador.
On 21 July 1978 I observed an adult Lesser Black-
backed Gull (Larus fuscus) from the CCGS Narwhal
at 57°11’N, 59°20’W approximately 140 km ENE of
Nain in the Labrador Sea. Sightings of Lesser Black-
backed Gulls include Greenland: Godthadb (ca.
64°30’N) and Godhavn (69°20’W) (Salomonsen
1967); at sea 100 to 675 km SSE of Cape Farewell,
Greenland (ca. 54°30’N to 60°00’N) (Brown 1968):
Northwest Territories (cf., American Birds 1978, 32:
1186; Alsop and Jones 1973); Churchill, Manitoba
(Ross and Cooke 1969): Grand Banks (Brown et al.
1975); Nova Scotia (cf., Nova Scotia Bird Society
Newsletter 1977, 19: 100; E. L. Mills, unpublished
data); and northeastern United States (W.R.
Petersen, unpublished manuscript; Bull 1964; K. D.
Powers, unpublished data).
The gull approached the ship from astern at 16:40
EST and “ship followed” for 55 min before departing.
It did not appear to be associated with any other
species in the area. The winds were WNW at 25 knots,
and seas were 1-2 m. The skies were overcast with no
precipitation. The bird was observed with 8 X 40
binoculars in good light within 50 m of the ship. All
distinguishing field marks of the species were seen,
446
including the bright yellow legs and feet. The mantle
was slate-gray becoming black in the outermost
primaries, suggesting L. f. graellsii. A color photo-
graph was obtained showing the mantle color pattern
of the bird.
The sighting is the first record of Lesser Black-
backed Gull for Labrador waters, and it adds to
evidence of increasing regularity off eastern North
America of the subspecies L. f. graellsii, which breeds
in Iceland, Faeroe Islands, the British Isles, and
Brittany (Witherby et al. 1941). The new record is only
about 2000 km distant from Iceland, well within the
range of sightings along American coasts from Davis
Strait to Cape Hatteras.
I express my appreciation to R.G. B. Brown,
W.R. Petersen, and T. Lloyd-Evans who helped to
improve this note. R. G. B. Brown of the Canadian
Wildlife Service arranged my passage aboard the
CCGS Narwhal, and traveling expenses. I am also
grateful to the United States Department of Energy
(DOE Contract No. EE-78-S-02-4706) for support.
Occurrences of the Red Phalarope
Adjacent States!
J. PAUL GOOSSEN2 and DANIEL G. BUSBY3
THE CANADIAN FIELD-NATURALIST
Vol. 93
Literature Cited
Alsop, J. F. and E.T. Jones. 1973. The Lesser Black-
backed Gull in the Canadian Arctic. Canadian Field-
Naturalist 87: 61-62.
Brown, R.G.B. 1968. Sea birds in Newfoundland and
Greenland waters, April-May 1966. Canadian Field-
Naturalist 82: 88-102.
Brown, R. G. B., D. N. Nettleship, P. Germain, C. E. Tull,
and T. Davis. 1975. Atlas of eastern Canadian seabirds.
Canadian Wildlife Service, Information Canada, Ottawa.
220 pp.
Bull, J. 1964. Birds of the New York area. Harper & Row,
New York.
Ross, R. K.and F. Cooke. 1969. Lesser Black-backed Gull
at Churchill, Manitoba. A new bird for Canada. Canadian
Field-Naturalist 83: 399.
Salomonsen, F. 1967. Fuglene pa grénland. Copenhagen.
[Bo QUT.
Witherby, H. F., F. C. R. Jourdain, N. F. Ticehurst, and
B. W. Tucker. 1941. The handbook of British birds.
Volume V. H. F. & G. Witherby Ltd., London. 356 pp.
Received 14 February 1979
Accepted 25 April 1979
in the Prairie Provinces and
Regent College, 2130 Wesbrook Mall, Vancouver, B.C. V6T 1W6
3Canadian Wildlife Service, P.O. Box 400, Fredericton, New Brunswick E3B 4Z9
Goossen, J. Paul and Daniel G. Busby. 1979. Occurrences of the Red Phalarope in the prairie provinces and adjacent
states. Canadian Field-Naturalist 93(4): 446-449.
Twenty-six records of Red Phalaropes ( Phalaropus fulicarius) were obtained for the prairie provinces and adjacent states. It is
suggested that the predominance of fall records in western Alberta and Montana is due to the guiding influence of the Rocky
Mountains for southbound and southwestbound birds, and that the scattered spring records result from the lack of such an
influence for northbound birds. It remains possible that the initial displacement from the normal migration route is caused by
adverse weather conditions.
Key Words: Red Phalarope, Phalaropus fulicarius, extralimital sightings.
Our interest in the Red Phalarope (Phalaropus
fulicarius) was aroused when we observed a female in
alternate plumage in the Assiniboine River Diversion
about 3km south of Lake Manitoba and 5 km
west of Delta, Manitoba (50°11’N, 98°19’W). This
was the third sighting in southern Manitoba; the
species is considered a rare spring migrant in the
Churchill region (Jehl and Smith 1970).
The Red Phalarope breeds along the coasts and
'This is paper number 74 of the University of Manitoba Field
Station (Delta Marsh).
islands of northern Asia, Europe, and North America.
It winters primarily in the southern hemisphere off the
coasts of South America and western Africa (Godfrey
1966). In North America, it migrates along both the
Atlantic and Pacific coasts (American Ornithologists’
Union 1957; Godfrey 1966). This note summarizes
and discusses Red Phalarope records from the
Canadian provinces of Manitoba, Saskatchewan, and
Alberta, and from the adjacent states of Montana,
North Dakota, and Minnesota.
Twenty-six records of the Red Phalarope in the
area under consideration are summarized in Table 1.
1979
NOTES
447
TABLE I—Red Phalarope sightings for the prairie provinces and adjacent north-central states
Location Date No. Source
Manitoba
East Shoal Lake 12 October 1963 | K. Gardner
Delta Marsh 19 June 1969 | R. E. Jones
Delta Marsh 30 June 1975 | This paper
Saskatchewan
Long Lake! July 1879 I Macoun and Macoun (1909)
Old Wives Lake? May 1895 l Macoun and Macoun (1909)
Sandfly Lake 11 June 1914 I Mitchell (1924)
Proctors Lake 21 May 1946 I| 22 Mowat (1946)
Alberta
Didsbury 3 September 1903 1 Salt and Wilk (1966)
Beaverhills Lake September 1925 I Salt and Wilk (1966)
North Saskatchewan River
near Graveyards Cabin 23 May 1953 4? Banfield (1954)
Pigeon Lake 13 July 1960 IPS Salt and Wilk (1966)
Carseland Dam? 23 October 1966 | Smith and Klauke (1967)
Cochrane Lake} 30 October 1966 | Smith and Klauke (1967)
Big Lake 11 November 1975 I Ebel (1976)
Stirling _ — Godfrey (1966)
Montana
Bowdoin National Wildlife Refuge July 1953 | P. D. Skaar
Harrison Lake 26 August 1959 | Rogers (1960)
Harrison Lake 3 November 1963 ] Rogers (1964)
Harrison Lake 11 October 1970 l Rogers (1971)
Freezeout Lake 17 August 1976 12 Serr (1977)
North Dakota
Cando4
Slade National Wildlife Refuge
Minnesota
Knife River Lake
Mille Lacs Lake
Mille Lacs Lake
Moorhead
23 May 1890
25 July 1963
17 November 1963
29 October 1976
19 November 1977
27 May 1977
'Now called Last Mountain Lake.
2Now called Johnstone Lake.
Stewart (1971)
Stewart (1971)
Hofslund (1964)
Savaloja (1977)
Harding (1978)
Wachtler and Wachtler (1977):
Anderson (1977)
3Validity of sighting questioned by Sadler and Myres (1976).
4Considered as hypothetical by Stewart (1971).
seven are of females; the others were not sexed. We
grouped the 25 observations with dates into three
periods: spring (April-June), midsummer (July), and
fall (August-November), and plotted them (Figure 1).
Two patterns are noticeable. First, seven of the eight
spring records and three of the four midsummer
records are scattered east of 110° W. Second, 9 of the
13 fall records are from the extreme western part of
the area, on or near the eastern slope of the Rocky
Mountains.
No correlation with weather could be established
for the Canadian sightings of Red Phalaropes and
P. D. Skaar (personal communication) noticed no
unusual, weather conditions associated with the
Montana sightings. The spring record for Minnesota,
however, was preceded by periods of thunderstorms
and unsettled weather with 35 to 40-km south winds.
The known eastern spring migration route follows
the Atlantic Ocean to the arctic breeding grounds.
Observations of Red Phalarope in southern Ontario
(W. E. Godfrey, personal communication), however,
suggest that some of these birds may pass through the
Great Lakes and Hudson Bay, using a similar short-
cut to that suggested for Arctic Terns (Sterna
paradisaea) (Godfrey 1973) and Sabine’s Gull (Xema
sabini) (Lambert 1973). The scattered spring migrants
reported on the Great Plains may be phalaropes
attempting to migrate overland. Oldsquaw (Clangula
hyemalis), also a northern breeder, regularly pass
through southern Manitoba in both spring and fall
(Sexton and Collins 1977), and the possibility of an
overland migration route for another arctic breeder,
448
FIGURE |. Red Phalarope sightings for the prairie provinces
and adjacent north-central states. Open circles, spring
records: half-filled circles, midsummer records:
closed circles, fall records.
the Long-tailed Jaeger (Stercorarius longicaudus),
has also been suggested (Jehl and Smith 1970).
The numerous fall records in Alberta and Montana
suggest that some migrants proceed southward or
southwestward until they encounter the Rocky
Mountains. With reference to the Montana observa-
tions, P. D. Skaar suggested that a prolonged Pacific
air flow might have deterred the birds from crossing
the mountain range (Rogers 1964), which thus served
as a leading line. No such guiding influence would be
available for northbound birds, which are much more
scattered. The fall sightings from Manitoba and
Minnesota may be migrants from the Hudson
Bay — Great Lakes route.
Two of the fall birds were listed as females in Table
1; however, we suggest that generally these are young
of the year having no previous migratory experience.
Of 18 casual records presented by Bent (1927) for
interior Canada and United States, 16 were fall
records and one was not dated. This further suggests
that dispersal of the young may account for many of the
interior records. It seems possible that adverse
weather conditions are the initial displacement factor
for these extralimital sightings, but that after dis-
placement the birds attempt the usual southward or
northward orientation, which leads some of them
overland.
Despite the periodic, infrequent sightings of Red
Phalaropes over the prairie provinces and adjacent
states, such records constitute an extremely small
fraction of the total migration of the species. It is
unlikely that enough of these birds survive for
THE CANADIAN FIELD-NATURALIST
Vol. 93
selection in favor of a regular inland migration to
occur.
Acknowledgments
We thank those individuals who replied to our
request for information regarding the Red Phalarope.
These include H. W. R. Copland, D. R. Crooke, J. B.
Falls, J. Ferrand, Jr., M. B. Fitzgerald, N. L. Ford, K.
Gardner, W. E. Godfrey, J. C. Green, E. B. Hazard,
J. Hines, P. B. Hofslund, E. O. Héhn,C. S. Houston,
R. D. James, R.E. Jones, F. McKanney. MEK:
McNicholl, W. J. Maher, D. Maurer, M. T. Myres,
R. A:-Paynter, P. D» Skaar, R: E-StewantsRewe
Storer, D. V. Weseloh, P. L. Wright, and R. L. Zusi.
We thank W. R. Salt and S. G. Sealy for reading an
earlier draft of the manuscript and also the reviewers,
J.B. Gollop and R.G.B. Brown, for their comments.
Literature Cited
American Ornithologists’ Union. 1957. Check-list of North
American birds. Fifth edition. Port City Press Inc.,
Baltimore, Maryland. 691 pp.
Anderson, E. 1977. Red Phalarope at Moorhead-Loon 49:
1973.
Banfield, A. W.F. 1954. Further notes on the birds of
Banff National Park, Alberta. Canadian Field-Naturalist
68: 182.
Bent, A. C. 1927. Life histories of North American shore-
birds. Part 1. United States National Museum Bulletin
142.
Ebel, G.R. A. 1976. Sightings of a Red Phalarope near
St. Albert, Alberta. Alberta Naturalist 6: 2.
Godfrey, W.E. 1966. The birds of Canada. National
Museum of Canada Bulletin 203: 1-428.
Godfrey, W. E. 1973. A possible short-cut spring migra-
tion route of the Arctic Tern to James Bay. Canada.
Canadian Field-Naturalist 87: 51-52.
Harding, H. 1978. Red Phalarope—Mille Lacs Lake. Loon
50: 45-46.
Hofslund, P. B. 1964. The Red Phalarope in Minnesota.
Loon 36: 25.
Jehl, J. R. and B. A. Smith. 1970. Birds of the Churchill
region, Manitoba. Manitoba Museum of Manand Nature
Special Publication |. 87 pp.
Lambert, K. 1973. The migration of Sabine’s Gulls. Xema
sabini, in the Northwest Atlantic. Canadian Field-
Naturalist 87: 57-60.
Macoun, J. and J. Macoun. 1909. Catalogue of Canadian
birds. Canada Department of Mines, Geological Surveys
Branch, Ottawa. 761 pp.
Mitchell, H. H. 1924. Birds of Saskatchewan. Canadian
Field-Naturalist 38: 101-118.
Mowat, F. 1946. Bird watching from a “jeep.” Blue Jay 4:
39.
Rogers, T. 1960. Northern Rocky Mountain — Intermoun-
tain Region. Audubon Field Notes 14: 56-58.
Rogers, T. H. 1964. Northern Rocky Mountain Region.
Audubon Field Notes 18: 57-60.
Rogers, T. H. 1971. Northern Rocky Mountain — Inter-
mountain Region. American Birds 25: 80-84.
1979
Sadler, T.S. and M. T. Myres. 1976. Alberta birds 1961-—
1970 with particular reference to migration. Provincial
Museum of Alberta Natural History Occasional Paper
No. 1.
Salt, W.R. and A. L. Wilk. 1966. The birds of Alberta.
Second edition. Queens Printer, Edmonton, Alberta.
511 pp.
Savaloja, T. 1977. Minnesota’s second Red Phalarope.
Loon 49: 44-45.
Serr, E. M. 1977. Northern Great Plains. American Birds
31: 190-194.
NOTES
449
Sexton, D. A. and K.M. Collins. 1977. Records of the
Oldsquaw in southern Manitoba. Blue Jay 35: 96-99.
Smith, W. and R. Klauke. 1967. A sight record of the Red
Phalarope in Alberta. Blue Jay 25: 25-26.
Stewart, R. E. 1971. Check list of birds of North Dakota.
Prairie Naturalist 3: 3-12.
Wachtler, D. and G. Wachtler. 1977. Red Phalarope at
Moorhead. Loon 49: 172.
Received 6 April 1978
Accepted 24 May 1979
Nesting of the Calliope Hummingbird in Kananaskis
Provincial Park, Alberta
DANIEL F. BRUNTON,! SIDNEY ANDREWS.,2 and DAVID G. PATON3
‘Southwick Drive, R.R. 3, Manotick, Ontario KOA 2N0
2General Delivery, Holland Landing, Ontario LOG 1HO
31481 Jalna Avenue, Mississauga, Ontario LSJ 1S6
Brunton, D. F.. S. Andrews, and D.G. Paton. 1979. Nesting of the Calliope Hummingbird in Kananaskis Provincial
Park, Alberta. Canadian Field-Naturalist 93(4): 449-451.
A nest of Calliope Hummingbirds (Ste/lu/a calliope) was studied in Kananaskis Provincial Park, Alberta, in 1977 and 1978.
Incubation period (16 d) and nestling period (18-21 d) were determined. The nest was constructed in precisely the same
location in successive years and nest construction was completed during incubation. These unusual nesting characteristics
may be adaptations to severe environmental constraints at high elevations near the northern limit of the species’ range.
Key Words: Calliope Hummingbird, Stellula calliope, incubation period, nestling period, nest construction. nest site
selection, Alberta.
A nest of the Calliope Hummingbird (Sre//ula
calliope) was discovered in Kananaskis Provincial
Park, Alberta, in mid-June 1977. This bird is found
only locally in Alberta (Salt and Salt 1976). A second
nesting was discovered at the same site in 1978. The
following discussion deals with the results of our
observations for both years.
Observations
The nest was situated on a horizontal limb of a
Lodgepole Pine (Pinus contorta) at the base of a steep
shale creek bank. It was | km south of the “Church
Camp” on the east bank of Pocaterra Creek (50°41’N,
115°5’W) at an elevation of 1660 m. The nest was a
tiny cup constructed of lichen fragments and fine
plant materials.
The 1978 nest was constructed on the weathered
remains of the 1977 nest: beneath the latter were the
remains of one or more previous nests (Figure |). In
1978 the nest was first observed on 10 June as the first
egg was being laid. The nest at that time was a shallow
cup with poorly defined walls and an unlined bottom.
At 11:45 the bird was observed frequently poking and
prodding the edges of the nest and pulling material in
towards the center. For about 2 min it performed an
action we described as “running on the spot” — ap-
parently deepening the cup in preparation for egg-
laying. The bird subsequently became very still on the
nest for several minutes and then flew off. A single.
pinkish-white translucent egg was subsequently found
in the nest. We believe the egg was laid as we were
watching the bird. On 13 June the nest contained one
dull-white, opaque egg and one (freshly laid) pinkish-
white, translucent egg. Although the first egg was laid
on the bare floor of the nest, both were now nestled
into a bed of feathers in a deep high-walled cup. The
increased size and depth of the nest was quite striking.
By 29 June the nest was larger yet, with substantially
thickened wall. The two-egg clutch of the 1978 nest
was completed on 13 June when we observed it at
16:20. On 29 June at 17:30 two tiny young were
observed, one still wet and lying on fragments of the
egg from which it had just hatched. The other young
was dry. Incubation had taken 16d. The nestling
period ran from 3 July to 21 July(18 d) in 1977 and 29
June to 20 July (21 d) in 1978.
On 20 July 1978 we observed the departure of the
young from the nest. One of the young was out of the
nest but still in the nest tree when observations began.
In approximately 20 min, the remaining nestling
FiGure 1. Adult (female) Calliope Hummingbird on nest in
Kananaskis Park. Alberta. Note remains of previous
nest under active nest: the 1978 nest was situated on
the remains of this one. Photographed on 23 June
1977 by D. G. Paton.
moved to the lip of the nest, made its first flight, and
ultimately moved 4m away. The female adult
returned to the nest tree four times to feed or preen the
young birds during the observation period.
The adult (only the female was seen in both years)
usually approached the nest from the creek side of the
tree, perching near the top of the nest tree and then
working down through it in three or four short flights.
The bird used the same perches each time. Calliope
Hummingbirds also exhibited a set pattern of man-
oeuvers to approach a feeder at Bow Valley Pro-
vincial Park, Alberta, and when disturbed the birds
would flee to precisely the same perch in a nearby
Lodgepole Pine (B. Romanyshyn, personal com-
munication).
During the early stages of incubation (up to
approximately 10 to 12 d), the adult would leave the
nest when observers approached within 10 m. During
late incubation and when young were in the nest,
however, the adult would allow the observers to
approach within | m of the nest before flying off.
THE CANADIAN FIELD-NATURALIST
Viole 93
When it was flushed during this latter period, the bird
would aggressively “buzz” the observers (usually
positioned 3 to 5 m from the nest). The bird would
often approach to within 30 to 50 cm of the observers
during this aggressive behavior.
Discussion
There is no published literature on the nesting of
Calliope Hummingbird in Canada with which we can
compare our observations. Calder (1971) cites 21 to
23 d for the nestling period in Wyoming and this
compares well to a figure of 21 d for a nest at Hood
River, Oregon (on file with the North American Nest
Record Scheme).
Figure 2 indicates the timing of egg and young
observations from cards on file with the British
Columbia Nest Record Scheme. None of the British
Columbia cards recorded a complete incubation or
nestling period observation. By summing the total of
days in which young or eggs were in these nests and
placing these data in quarter-month periods, the
nesting chronology of the Calliope Hummingbird in
British Columbia is illustrated. The total nesting
period is roughly 37 d. The difference between starts
of young and egg peaks is about 15 d. The dates for
Kananaskis Park in 1977 and 1978 agree well with the
peak of observations of eggs and young in nests in
British Columbia. The sample of British Columbia
records was too small to subdivide by latitude and/or
elevation. When sufficient numbers of observers are
filed, this would certainly be worthwhile.
The total length of the nesting period for Calliope
Hummingbird is approximately 34 to 38 d in north-
western North America (Calder 1971: this study).
There is some suggestion that the period is shorter at
the northern limit of the species range (34-37 d in
Kananaskis vs. 36-38 d in Wyoming).
We strongly suspect that the same female was
responsible for the 1977 and 1978 nestings in
Kananaskis Park. As the bird was not banded, we
cannot be certain of this. In any case, the use of
precisely the same site in successive years 1s apparently
atypical in hummingbirds. W. Ray Salt (personal
communication) observed this practice with Rufous
Hummingbirds (Selasphorus rufus) in the 1960s in
British Columbia, and has a nest collected by J. Grant
of Vernon, British Columbia, which was used in 1938
and again in 1939. Pearson (1953) notes repeated use
of nests by the Estella Hummingbird ( Oreotrochilus
estella) in high mountain caves in the Peruvian Andes.
He indicates, however, that these nests are “recon-
ditioned” each year: the Calliope Hummingbird nests
we observed were completely rebuilt upon the
previous year’s remains. We found no published
reports of nests being completed during the period of
egg laying, although Skutch (1951) does report
1979
451
Nestin
NOTES
25
20
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6 :
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5 i | 6
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£ 10 HE +
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: \
5 \
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\
e
OP Pcccecce,
June
( quarter
July
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FIGURE 2. Frequency of occurrence of eggs and young in nests of the Calliope Hummingbird in British Columbia.
renewal of some nesting material during incubation
by some tropical hummingbirds.
The utilization of the same nest site for a succession
of years, completion of nest construction during
incubation, and the possibly shorter nesting period of
the species in this area may be adaptations to near-
marginal breeding conditions in Alberta at the
northern limit of its range.
Acknowledgments
Field assistance and additional data were provided
by C. Backer, A. Masters, and B. Romanyshyn.
H. W. R. Copland and J. B. Gollop provided nesting
data from the Prairie Nest Record Scheme, as did
R. W. Campbell from the British Columbia Nest
Record Scheme and J. Crump from the North
American Nest Record Card Program. R. D. Strick-
land contributed an extensive collection of humming-
bird literature. W. R. Salt provided unpublished data.
and observations from earlier studies. Several earlier
drafts of the manuscript were typed by B. Wackerle.
Our thanks to all these people for their valuable
assistance.
Literature Cited
Calder, W. A. 1971. Temperature relationships and nesting
of the Calliope Hummingbird. Condor 73: 314-321.
Pearson, O. P. 1953. Use of caves by hummingbirds and
other species at high altitudes in Peru. Condor 55:
17-20.
Salt, W.R. and J. R. Salt. 1976. The birds of Alberta.
Hurtig Press, Edmonton.
Skutch, A. F. 1951. Life history of Longuemare’s Hermit
Hummingbird. Ibis 93: 180-195.
Received 24 February 1978
Accepted 3 April 1979
News and Comment
FON Conservation Award for The Canadian Field-Naturalist
A Federation of Ontario Naturalists (FON) Con-
servation Award has been presented to The Ottawa
Field-Naturalists’ Club in recognition of its out-
standing contribution to the cause of conservation.
The citation for the FON Award read as follows:
“for the continued excellence of the journal
The Canadian Field- Naturalist which has a well
deserved national and international reputation
and which provides a vital forum for reports on
natural history and the environment by ama-
teur and professional observers alike.”
The FON Conservation Award was accepted by Dr.
Lorraine C. Smith, Editor of The Canadian Field-
Naturatist, at the Federation of Ontario Naturalists
Conference at Toronto, Ontario on 5 May 1979. Dr.
Peter A. Peach, President of the FON expressed the
appreciation of all FON members for the continued
excellence of the journal and personally congratulated
the Editor. Mike Singleton, General Manager of the
FON, added his congratulations and commendation
to those of the Awards Committee, FON Directors,
and participants at the annual conference and pointed
out that the award in recognition of The Canadian
Field- Naturalist was well earned. Moreover, he wrote:
“It seems to me that we do not often enough recognize
the great contribution the Ottawa club has made
through this journal in facilitating natural history
reports by amateur and professional alike, and most
importantly in stimulating research by amateurs
across the country.” He asked that the commendation
and congratulations be drawn to the particular
attention of all members of The Ottawa Field-
Naturalists’ Club. °
Notice of The Ottawa Field-Naturalists’ Club Annual Business Meeting
The 101st Annual Business Meeting of The Ottawa Field-Naturalists’ Club will be held in the auditorium of
the National Museum of Natural Sciences, Metcalfe and MacLeod, on Tuesday, 15 January 1980, at 8:00 p.m.
452
Diana R. Laubitz,
Recording Secretary
Book Reviews
ZOOLOGY
Guide to the Study of Animal Populations
By James T. Tanner. 1978. University of Tennessee Press,
Knoxville. 186 pp. U.S. $8.95.
Many of the mathematical and statistical tech-
niques applied to the study of animal populations
elude all but a specialist in the field. Therefore, I was
somewhat elated to discover that this book is
comprehensible to an average biologist having some
experience in biometrics.
The book contains ten chapters describing a variety
of techniques for studying animal populations. Sub-
jects such as Density and Dispersion, Mark-Re-
capture Methods, Sex and Age Composition,
Mortality and Survival, Reproduction, Migration
and Recruitment are presented. Considerable
discussion is devoted to mathematical relationships
between population characteristics and methods of
predicting population changes.
As the title suggests, all examples given deal with
animal populations and many deal with big game
species. Several problems are worked out in each
chapter and suggestions are given for the arrangement
of the raw data. These examples do much to dispel the
mystique of seemingly incomprehensible formulae.
Symbols are used as consistently as_ possible
throughout the text and a list is provided at the
beginning of the book.
The book is replete with references to more detailed
discussions of various subjects and methodologies
available in the literature. Moreover, the text is
conveniently cross-referenced from chapter to
chapter, enabling the reader to refresh his memory
quickly from previous presentations on a topic.
One of the most interesting chapters discusses
mark-recapture methods for measuring population
characteristics. This is a very practical presentation
and provides valuable insight into research design
compatible with statistical analysis of the data.
The publication is intended for use by graduate
students and field biologists. I am certain that it could
be a very useful addition to any biological bookshelf.
The straightforward presentation and numerous
examples bolster the reader’s confidence that he can
apply statistical methodologies and _ techniques
competently and that studies of animal populations
need not bea mathematical maze. The author’s candid
critiques of techniques are valuable. Also, the list of
literature citations is a useful tool for non-specialists
wishing. to delve further into some aspect of
population studies.
DAN MURPHY
Wildlife Service, Government of the Northwest Territories,
Yellowknife XOE 1HO
Quaternary Vertebrate Faunas of Canada and Alaska and Their Suggested Chronological Sequence
By C. R. Harington. 1978. Syllogeus Number 15. National
Museums of Canada, Ottawa. 105 pp., 15 illus. Available
from Library of National Museum of Natural Science,
Ottawa KIA OMS8.
Most natural historians working with recent faunas
are not especially familiar with the literature dealing
with Pleistocene faunas and natural history. This is
understandable, considering the present volume of
literature dealing strictly with modern faunas. Any-
one working with modern groups, however, can
appreciate the historical perspective provided by the
paleontological literature, especially that of the
Pleistocene. Often this historical perspective provides
insight into modern animal and plant distributions
and geographic patterns as well as an understanding
of changes in their geography in response to changes
in ancient drainage patterns or patterns of glaciation.
A good synoptic review serves two functions. First,
it provides a broad generalized background on a
subject. Secondly it introduces the nonspecialist to the
literature dealing with that subject. With regard to the
Pleistocene faunas of Canada, Harington’s paper
successfully fulfills both objectives.
The information is presented in a concise, straight-
forward manner that makes data retrieval simple. The
faunas are listed for each province and Alaska. Each
section opens with a narrative concerning various
fossil discoveries and their significance with regard to
understanding Pleistocene events in the province.
Both marine and terrestrial vertebrates are described.
There is a good balance in the text between
descriptions of the history of collecting, outstanding
specimens that have been found, and the type of
information that these finds provide. Following the
453
454
opening narrative of each section the Pleistocene
faunas of specific provincial localities are listed. It is at
this point that the paper becomes a reference work.
Each faunal description begins with the fauna’s
location. The locality information varies in detail
from the very general such as “Hand Hills, Alberta
near Delia,” to the very specific: “Acasta Lake,
Northwest Territories (65°24’N, 115°31’W).” The
locality description is followed by a faunal list which
includes all known vertebrates from the site. This part
alone will be of use to anyone interested in determin-
ing the Pleistocene distribution of a particular species.
The mammals are well covered; the birds, reptiles,
amphibians and fish are not as complete, but this isa
reflection of the state of the art and the availability of
specimens. It points out sharply the areas where more
research is needed. Each faunal list is followed by a
suggested age for the locality. Radiocarbon dates are
included when available. The last part of each faunal
description, entitled “Remarks” includes a diverse
amount of information, such as descriptions of the
bone deposits, environment of deposition, interpre-
tation of habitat and the first occurrences of particular
species. Finally, there is a listing of references for that
particular fauna.
The paper has sixteen illustrations. One is a map
showing the location of all the faunas discussed, one 1s
a chart with the suggested chronological sequence of
Birds of Man’s World
By Derek Goodwin. 1978. British Museum (Natural
History) and Cornell University Press, London and
Ithaca. villi + 193 pp., illus. U.S. $10.95.
Although the title may suggest to some another
“gloom and doom” volume on Man’s destruction of
wildlife, the subject matter concerns the ability and
inability of birds to adjust to human environmental
changes. Through a series of examples, Goodwin
illustrates how the natural habits of various species
suit them for adapting to, and thriving in Man’s
world, while those of other species doom them to
extinction. This book appears to be written primarily
for the layman, but as pointed out by D. W. Snow in
the foreword, it also contains much of value to the
professional ornithologist.
The majority of Goodwin’s examples are European,
with considerable emphasis on personal experience;
however, other continents are not neglected, with
many examples coming from the literature. Unfortun-
ately; the sources are rarely given, and then usually
vaguely. Thus, the serious student who wishes to
pursue a topic in greater depth has little help in
THE CANADIAN FIELD-NATURALIST
Vol593
the faunas, three are artistic reconstructions of
specific Pleistocene vertebrates, and eleven are photo-
graphs of specimens. All of the photographs occupy a
full page. The map could have been a little larger as the
amount of reduction tends to make reading the
numbers of the localities difficult, but with a little
squinting they can be deduced.
The final discussion and summary places the entire
history of the Pleistocene of Canada in perspective. It
discusses chronologically first appearances, migra-
tions, changing environments and their significances
with regard to changes in the fauna. The reference
section has plenty of citations, which will permit
anyone interested to pursue a particular subject
further. The appendix contains a listing of nine faunas
from northeastern Siberia for comparison with those
from Canada and Alaska. This section is of interest
since the faunal lists include species that did not
manage to cross the Bering Straits into North
America.
This well-ordered, concise review will makea handy
starting point for anyone who wishes to obtain an
overall view of the Pleistocene in Canada and Alaska.
H. GREGORY MCDONALD
Department of Vertebrate Palaeontology, Royal Ontario
Museum, Toronto, Ontario M5S 2C6
locating original material.
In his introduction, Goodwin points out that his
book is not intended as a thorough review covering all
aspects of Man’s interactions with birds. The six
numbered chapters cover various aspects of this topic
as outlined below. The text is chatty and non-
quantitative but written well and apparently free of
printing errors. Several photographs and line illustra-
tions by Robin Prytherch enhance the book, although
neither are of exceptional quality.
Chapter |, “Man and the environment,” is concern-
ed with the effects on birds of habitat alteration. These
include obvious direct effects, such as destruction of
habitat for some species, and expansion of habitat for
others. Indirect effects are also discussed. For
example, although many species benefit from the
provision of artificial nest sites, the increase in
aggressive hole-nesters such as Starlings could result
in a decrease in less aggressive hole-nesting species.
Although not intended as a complete review, this
chapter seems comprehensive, with a few minor
omissions. A long discussion about birds coming to
1979
drink at artificial watering-holes in desert areas might
well have mentioned the large number of recent
occurrences of marine birds far inland at large
reservoirs in the southern USA. I found Goodwin’s
discussion of Purple Martins and Tree Swallows
misleading. After mentioning the benefit to the
martins of numerous houses built for them, Goodwin
notes that Tree Swallows have also been incidental
beneficiaries by using nest boxes which have not
attracted martins. This is true, but surely the Tree
Swallow has benefitted far more from the extensive
bluebird house programs in the prairie provinces and
other parts of North America. The appearance of
hybird bluebirds and swallows during range expan-
sions caused by the provision of artificial sites is also
omitted.
Chapter 2, “Birds in towns,” consists of a compari-
son of major cities in various parts of the world in
relation to their avifauna. Interactions of human
behavior and bird behavior are shown to determine
both the extent birds will live in cities and which
species will do so under various circumstances.
Chapter 3, “Birds fed by Man,” includes discussion
of birds as pests, birds as scavengers on garbage, and
deliberate feeding of birds by people. As in other
chapters, Goodwin emphasizes the natural behavior
of species as it adapts them to respond to food
provided by Man intentionally or otherwise. Missing
from the discussion of birds in crops are cases of birds
saving crops, such as California Gulls in Utah and
Franklin’s Gulls in Manitoba. This chapter overlaps
to some extend both chapters 2 and 5.
Introduced birds on all continents form the subject
BOOK REVIEWS
455
matter of chapter 4. After pointing out that most
species that do well on being introduced into a new
place are species that did well in their places of origin,
where they were frequently somewhat of a pest,
Goodwin reviews the success and failure of various
introductions, and comments on factors which may
determine rate of success. This chapter is slightly
outdated for North America: the House Finch now
inhabits much more of the East than indicated: the
Skylark has spread to the San Juan islands of
Washington (at least temporarily), and the predicted
action to control the Monk Parakeet in the USA has
taken place. I was also surprised to find only brief
mention of the Black Swan of New Zealand with no
reference to its adverse effects on native waterfowl
there.
In chapter 5, Goodwin compares several species as
to their habit of eating an unnatural food — bread. He
ponders how each might come to recognize bread as
food, howeach behaves in obtaining and eating bread,
and how much competition for bread among species
takes place.
In his final chapter, Goodwin offers a cautiously
optimistic outlook on the future. A list of scientific
names of birds discussed includes page numbers, and
thus also serves as a species index.
I recommend this book as a highly readable essay
on an important topic.
MARTIN K. MCNICHOLL
Beak Consultants Ltd., 3530 11 A St. N.E., Calgary, Alberta
T2E 6M7
Statistical Inference from Band Recovery Data: a handbook
By C. Brownie, D. R. Anderson, K. P. Burnham, and D. S.
Robson. 1978. Resource Publication Number 131, U.S.
Fish and Wildlife Service, Washington, D.C. No price
given.
Many workers with fish, mammals and birds have
need of mark-recapture estimation techniques for
assessment of population sizes and mortality rates. It
is not that better estimates are provided by such
techniques, but rather that direct sampling of
concealed and highly mobile populations is
impossible. Given the need for them, mark-recapture
methods have evolved to levels of sophistication that
would greatly impress early workers in the area. This
handbook represents the latest state of the art,
reflecting several decades of improvements in two
areas: precision and accuracy.
Students encountering the simplest mark-recapture
methods for the first time tend to react with shock at
the low precision (i.e., wide confidence intervals) of
their estimates, and often are convinced that they
must have made a mistake in their calculations. The
methods also tend to be vulnerable to biases in
estimation caused by failure of assumptions, and such
biases result in low accuracy. Methods such as those
presented in this book improve precision by using
data from repeated marking and capturing overa long
series of time intervals (usually years), and they deal
with biases by presenting a hierarchical series of
models which differ from each other in their
assumptions about the parameters of the populations
being estimated. The authors emphasize that results
are still dependent on assumptions for any particular
model, but this approach allows sequential testing of
the validity of various assumptions, which leads to
456
selection of the model satisfying the conditions with
which the researcher is dealing.
Although the authors hope they have produced “a
simple, easy-to-read primer.” have emphasized
examples from real data, and have put some of the
mathematics in appendices, this handbook is not for
the beginner. Differential calculus and two or three
courses in applied statistics are assumed, and anyone
without prior knowledge of the basic principles of
mark-recapture methods is advised to start with the
treatment of this subject in a general quantitative
ecology text. For the professional working on wild
populations that can not be assessed directly,
however, this handbook will be a necessary addition
to his library. The discussion and examples are in
terms of bird-banding studies, but the models are
generally applicable. Equal time intervals and
recovery of dead animals (marked or unmarked) are
assumed, with some discussion of unequal intervals
and of capture and release of live animals. All
methods described are related to FORTRAN
computer programs designed for IBM systems, with
The Gannet
By Bryan Nelson. 1978. Buteo Books, Vermillion, South
Dakota. 336 pp., illus. U.S. $25.
It is obvious that Nelson has spent many hours
watching Gannets. His readable style combines dry
and factual with informal and anecdotal accounts. An
incredible amount of detail is found throughout the
book with emphasis placed on fact, integration, and
interpretation. Nelson assures the reader that this
book is not an extract of his recent monograph (The
Sulidae: Gannets and Boobies. 1978. Oxford Uni-
versity Press. 1012 pp. U.S. $98) but has been written
afresh and with a different approach.
The book is organized into seven chapters. Addi-
tional introductory and final sections include a
preface, acknowledgments, a thumbnail sketch intro-
duction, two pages on names, references, bibliog-
raphies, and an index. The 32 tables are grouped
together and follow chapter 7. Major points are set
out in numbered statements in a summary for each
chapter. Numerous illustrations by John Busby are
scattered throughout the book and are particularly
valuable in the behavior chapter; a two and one-half
page schematic representation of the Gannet’s life
history is especially nice. Black-and-white photo-
graphs covering 32 pages are located in the middle of
the book. Although figures are numbered, the text
often does not refer to them. The map numbers
corresponding to the locations of the six Canadian
THE CANADIAN FIELD-NATURALIST
Vol. 93
versions converted for use on CDC systems also
available. Detailed instructions for data coding are
given, and small jobs can be sent (as punched data
card decks) to the Patuxent Wildlife Research Center
in Maryland for analysis and commentary at no
charge. The loan of tapes can be requested by those
wishing to obtain the programs, and they will be sent
without charge to be copied and returned.
The authors strongly emphasize the need for
advance planning of any study based on a thorough
understanding of the methods and their assumptions.
Chapter 8 summarizes analysis methods proposed by
previous workers, and discusses interpretation of
mark-recapture analysis results. Chapter 9 considers
the planning of a study, including timing of sampling
and necessary sample size. There is a comprehensive
bibliography.
R. H. GREEN
Department of Zoology, University of Western Ontario,
London, Ontario N6A 5B7
gannetries in Figure 10 also do not correspond to the
numbers in the text. In general a better correlation
between text citations and figures, tables, and illustra-
tions would have been an improvement.
Over 100 pages are devoted to breeding behavior
and ecology where the author presents a myriad of
interesting facts and ideas. An interpretation of the
function and survival value of various behavioral
traits 1s usually provided. The numbers and distribu-
tion chapter contains excellent sections on how, when,
and what to count. Another interesting section
discusses fishing techniques and includes quotes from
Scottish fishermen.
Chapter 6 on the Gannet family and the order seems
to come too late in the book. Chapter 7, on the other
hand, is a perfect final chapter with subchapter
headings on such fascinating topics as early accounts
of the Gannet’s natural history, its use as food, man’s
practice of culling in breeding colonies, oiled birds, its
behavior in captivity, and its role in art and literature.
I believe that those who read the book will find
immense enjoyment. Buteo Books should be com-
mended for publishing and making available several
important avian works originally published in Europe.
NOEL J. CUTRIGHT
Wisconsin Electric Power Co., 231 W. Michigan, Mil-
waukee, Wisconsin 53201
1979
BOTANY
Vascular Plant Families
By James Payne Smith, Jr. 1977. Mad River Press, Inc.,
Route 1, Box 151-B, Eureka, California 95501. 320 pp.,
illus. $9.55.
Students and teachers of plant taxonomy alike will
welcome this elementary book which contains every-
thing from an outline of the taxonomic hierarchy and
scientific names, through family descriptions, lists of
regional floras for the United States, notes on how to
collect plants, to a useful glossary.
Following the system of Cronquist, Takhtajan, and
Zimmerman (1966) for divisions, classes, subclasses
and orders, and Cronquist (1968) for the sequence of
families, the author presents “an introduction to the
families of vascular plants native to North America,
and selected families of ornamental or economic
importance.” This introduction consists of short
descriptions of the families, an indication of the
number of genera and species, a few selected genera,
BOOK REVIEWS
457
some recognition characters, and a floral formula.
Over one hundred plates of fine line drawings by
Kathy Simpson accompany this text. Of special
interest to the student is the chapter entitled “Vege-
tative morphology of the flowering plants.” This is an
illustrated glossary which is divided into sections that
give the terminology used to describe various parts of
a plant and their arrangement, e.g., roots, stems,
leaves, etc.
The book is written for students by a teacher who
has tried to present his subject as simply as possible.
The elimination of all but a few of the non-North
American families has certainly made this easier. The
author is to be congratulated on a job well done!
WILLIAM J. CODY
Biosystematics Research Institute, Canada Agriculture,
Ottawa, Ontario KIA 0C6
An Atlas of Airborne Pollen Grains and Common Spores of Canada
By I. John Bassett, Clifford W. Crompton, and John A.
Parmelee. 1978. Agriculture Canada, Ottawa. 350 pp.,
illus. $12 in Canada; $14.40 elsewhere.
This attractively and robustly bound research
monograph represents the culmination of over two
decades of careful, well reported investigations by the
authors into the problems of airborne allergenic
pollen and spores. It isaddressed to workers who wish
to identify airborne pollen and spores because of their
interest in either allergy problems, or in general and
applied palynology. Its effectiveness therefore should
be measured by the clarity and quality of the
illustrations; the rigor and decisiveness of the keys:
and the degree to which it will become indispensable
to the laboratory scientist in the appropriate dis-
ciplines. But its format implies a wider readership
target.
The first 39 pages introduce the subject of allergy
pollen. They vary from a few fascinating pages
(15-16), to some dreary tables (16-39), which should
be appendices. Pollen grains are exciting objects. A
lively up-to-date chapter on their development, role in
reproduction, life-cycle strategies, paleoenvironmen-
tal reconstructions, etc., as well as a more animated
discussion of the intriguing material on p. 13 on
diurnal patterns and the spectacular data on the
decline of ragweed fallout since the 1950s might have
captured a wider readership. And what about allergic
reactions to pollen? Couldn’t we have had a few pages
summarizing the state of the medical arts on that?
We are plunged into the dichotomous identification
keys to pollen, the heart of the book for the user.
Presumably, the user will be the technician in the
allergy laboratory, because other pollen work deals
with ranges of material incompletely covered in this
book.
Pollen keys at best are unsatisfactory and they are
usually fully comprehensible and usable only by the
authors themselves. The keys in this book are useful
but uneven. For example, the distinctions between
important species (Fagus, Quercus) are not clearly
handled. Comptonia peregrina is not consistently
tetraporate, as Figure 107 C and D demonstrate, and
in fact is indistinguishable from Myrica. On the other
hand A/nus could be separated readily to species, if it
were useful to do so from the allergy viewpoint.
The photomicrographs are of very uneven quality.
Some taxa have no light microscope photographs
(Carex), others have interference and SEM but no
non-interference light microscope illustrations
though they are essential (e.g.. Fraxinus, Luzula,
Triglochin, Zea, and others). The magnification varies
widely from plate to plate. And no attempt has been
made to achieve uniformity of positioning of the
grains in the SEM, photographs essential if they are to
be used diagnostically. We are told that interference
458
photomicroscopy was employed because of its great
use in such groups as Urtica. Possibly, but it is far
from convincing that its use in most of the other
groups illustrated adds anything to the book.
Part 2 deals with fungus spores, but it differs
markedly from Part | in having no keys, in offering
only a selection of representative types and illustra-
tions, and in providing nothing more than a glossary
to initiate the novice into the distinctive descriptive
ENVIRONMENT
THE CANADIAN FIELD-NATURALIST
Vol. 93
terminology of mycology. It creates the impression of
being an afterthought.
This book will find its way into most North
American pollen-spore labs, but whether it becomes
dirty and dog-eared with use is another question.
J.C. RITCHIE
Division of Life Sciences, Scarborough College, University
of Toronto, West Hill, Ontario MIC 1A4
A Vanished World: the dinosaurs of western Canada
By Dale A. Russell. 1977. Natural History Series Number
4. National Museum of Natural Sciences, National
Museums of Canada, Ottawa. 142 pp. $12.95.
In his introduction to A Vanished World, Dale
Russell states that “The purpose of this volume is to
visualize, so far as possible, the vanished world of
Canadian dinosaurs . . . it is hoped that this work will
help others to understand what western Canada may
have been like during a very interesting and relatively
well documented period of its physical history.” Upon
reading the book, one cannot help but conclude that
in the short space of 142 pages, the author has
managed to manifest successfully a most original
approach to the study of the dinosaur era.
The book is filled with superb photographs (by
Susanne M. Swibold) of the Canadian badlands, and
of modern vegetation and habitats that compare
closely with the projected environments of the Late
Cretaceous in Canada. The photographs are com-
plemented by equally striking artistic reconstructions
(by Eleanor M. Kish) of many of the Canadian
dinosaurs described in the text. All photographs and
illustrations are in color.
The first chapter deals with the history of dinosaur
collecting in western Canada, and as most published
histories of palaeontology are concerned mainly or
only with American localities and collectors, much of
the material will be new to the general reader.
The second chapter provides a general survey of the
geology of the Canadian badlands, witha brief insight
into the methods of interpreting the prehistoric
geography of an area from its sediments.
The third chapter describes Dinosaur Provincial
Park in Alberta, an area of about 15000 acres of
badlands containing exposed sediments that are from
76 to 73 million years old. At that time, Dinosaur
Provincial Park was part of an ancient alluvial plain
situated between the Rocky Mountains and the great
inland sea which then covered much of the interior of
the North American continent. Considering in turn
the sediments themselves, then their plant, inverte-
brate, and vertebrate remains, the author logically
and precisely reconstructs the environment of that
period. A short section at the end of the chapter invites
the reader to imagine himself back in those times and
then to consider the possibility of his own survival
under such conditions.
The fourth chapter treats the badlands around
Drumheller, Alberta in a similar manner. The
sediments of the Drumheller region contain fossil
remains ranging from 72 to 70 million years in age. At
that time, Drumheller was located on the southern
edge of a large delta that extended more than 200
miles from the Rocky Mountains to the inland sea.
The delta was “an enormous low-lying wetland of
shallow lakes, swamps and marshes, traversed by
streams that fanned out from the major river courses
to the edges.” Various aspects of the ecology of the
delta are discussed, from the patterns of deposition of
sediments to the vegetation and the dinosaurs that
inhabited the area. Over a period of a few hundred
thousand years, the ecosystem changed and the delta
developed the characteristics of a coastal plain, as
stream beds shifted and the swamps drained. The
vegetation became markedly different from that of the
wetland period, with forests of broad-leaved trees,
many of which are familiar to us today. The kinds of
dinosaurs inhabiting this drier, hardwood-forested
area were somewhat different from those that lived in
the swamps.
The fifth chapter takes us through the sediments of
70 to 65 million years ago into the geological period
that signalled the end of the dinosaurs. The vegetation
and the dinosaurs that lived on the large subtropical
floodplain of this period are discussed. Because the
floodplain extended into the United States, con-
temporary dinosaurs from Montana, Wyoming, and
the Dakotas are also reviewed.
1979
The last chapter considers the extinction of the
dinosaurs. The final testimony of the late Cretaceous/
Early Paleocene rocks is discussed, and various
general aspects of the extinction problem are ex-
amined. In a concluding section, entitled “From the
Depths of Space,” the author takes the opportunity to
elaborate upon the extinction theory he himself
favors most: that of a cosmic event (probably a
supernova) that bombarded the Earth with “waves of
high-energy radiation comparable to those occurring
near powerful thermonuclear explosions.”
The bibliography, while comprehensive, is not
overwhelming to the non-scientist, and many of the
Biogeography and Ecology of Southern Africa
Edited by M. J. A. Werger. 1978. W. Junk, The Hague.
xvi + 1439 pp., illus. (in 2 volumes). DFL 365.
Southern Africa as referred to in these two volumes
covers an area of some 6100000 km?. The geo-
political areas included are: Angola, part of Shaba,
Zambia, Malawi, Mozambique, South West Africa,
Botswana, Rhodesia, South Africa, Swaziland, and
Lesotho. Southern Africa varies in altitude from sea
level to 3482 m and has a wide range of biotic and
abiotic features.
There is an extensive published and unpublished
literature on the biogeography and ecology of
southern Africa. Much of this material appears in
local journals, in various government reports, and as
theses. As such it is not readily available to interested
biogeographers and ecologists outside southern Afri-
ca. These volumes successfully review this large body
of knowledge. As Werger points out, the ecological
aspects covered are mainly of a descriptive nature,
concentrating on diversity and variation in species
and on a structural-functional evaluation; physio-
logical aspects of the ecology, and energy and nutrient
flow systems remain virtually untouched. Research in
these latter fields has only recently started in southern
Africa.
The two volumes are divided into 41 chapters.
Approximately 75 percent of the contributing authors
were living and working in South Africa when they
wrote their chapters. Chapters | to 9 cover the
physical and geological environment, past and pres-
ent. Chapters 8 to 14 each deal with a major botanical
biome while Chapters 16 to 31 each cover a major
zoological taxonomic group. The next nine chapters
deal with the biogeography and ecology of special
habitats such as lakes, rivers, marine habitats, high
termitaria, and heavy metal and other toxic soils. The
final chapter discusses the conservation of southern
BOOK REVIEWS
459
references are government publications which are
reasonably accessible to the general public.
A Vanished Werld is written in a clear and logical
style, and the ancient environments reconstructed
from the rocks and their fossil remains become strong
visual images within the mind of the reader. This book
will be a valuable addition to the library of any person
who has an interest in natural science.
JOANNE E. LINDSAY
Department of Vertebrate Paleontology, Royal Ontario
Museum, Toronto, Ontario MSS 2C6
African ecosystems. An extensive use of photographs
and illustrations is made throughout the text. Typo-
graphical errors were few and the largest single
printing mistake (Figure 9, Chapter 39) is corrected
with an erratum.
A major pitfall of such reviews can be the length of
time required to get them published. Although it is not
possible to eliminate completely this lag phase, the
publishers can do much to make it as short as possible,
thereby increasing the value of the review. Dr. W.
Junk, publishers, should be commended for the speed
in which these two volumes were brought to press.
References in the text date as recently as 1977.
Dr. Werger’s contribution to the volumes as editor
has been considerable. Throughout the text he has
constructed a network of cross-references to earlier or
later chapters in which additional information or a
difference of interpretation can be found.
Unfortunately, in many chapters readers are refer-
red to a specific geological formation or plant
community located near a town not likely to be known
beyond southern Africa or to individuals who have
never lived there. A much greater use of maps such as
Figure | in Chapters 30 and 38 would have been very
useful.
Readers who have only a lay interest in the
biogeography and ecology of southern Africa will find
general information on many subjects concerning a
most interesting part of the world. For example, in
Chapter 16 we are informed that the world’s largest
earthworms, measuring 7 m in length by 7.5 cm in
diameter, occur near Debe Nek in the Cape Province
in South Africa and that some savanna areas have
rodent populations that exceed those of the Brown
Lemming in the tundra of Alaska at cycle peaks
(Chapter 31).
460
These two volumes should attract many readers in
academic or research institutions with an interest in
Africa. Although, as Dr. Werger notes, the books are
not all-encompassing (both in subject or in literature
citations), they are an excellent source of information
for any amateur or professional ecologist, naturalist,
or biogeographer beginning a project on southern
Africa. Purchase of these volumes by interested
The Ecology of North America
By V.E. Shelford. 1978. University of Illinois Press, Ur-
bana. Paperback edition, originally published 1963. 610
pp., illus. No price given.
This book was first published in 1963, and is now
resurrected in paperback form. Undoubtedly an
important contribution to ecology in its time, the
purpose of the book is“. .. to describe North America
from an ecological viewpoint as it appeared in the
period A.D. 1500 to 1600 before European settle-
ment.”
Taking over half a century to complete, Shelford
undertook the monumental task of reconstructing the
ecological story of primeval North America using
results from his own studies, those of his students and
his colleagues. This book is designed to assist the
ecologist in understanding the habits, biotic com-
munities, and the distribution and abundance of
plants and animals in primeval North America. To
this end, Shelford focuses on the biogeographical
aspects of primeval ecology.
The text is organized into 19 chapters. The first
chapter reviews the meaning of major underlying
ecological principles. Among them are habitat,
community, dominance, community development,
population densities and distribution, life span, life
cycles, solar radiation, temperature and moisture,
natural communities of North America, and com-
munity terminology. Many of these principles con-
tinue to form the underlying components of modern
Chlorinated Phenoxy Acids and their Dioxins
Edited by C. Ramel. 1978. Ecological Bulletins Number 27,
Swedish Natural Sciences Research Council (NFR),
Stockholm, Sweden. 302 pp. 75 SwCr.
This bulletin is a collection of papers from a
conference arranged by the Royal Swedish Academy
of Sciences in Stockholm, 7-9 February 1977. The
intent of the symposium was to provide an up-to-date
summary of objective scientific data on chlorinated
phenoxy acids and their dioxins, and to make a risk-
benefit evaluation of this data in a series of work
THE CANADIAN FIELD-NATURALIST
Vol. 93
individuals, however, is unlikely since the approxi-
mate Canadian price of $200 will be prohibitive in
most cases.
RICHARD D. ROBARTS
Schultz International Limited, 1155 West Georgia Street,
Vancouver, British Columbia V6E 3H4
ecology; however, a decade and a half of research and
development has improved and expanded our knowl-
edge of them. In this regard it is important that the
reader be aware of these changes.
The following 18 chapters cover all of the major
communities in North America, from the high Arctic
to southern Florida and Cuba.
Shelford not only attempts to document the abiotic
and biotic components of each community, he
attempts to interrelate these components and develop
a comprehensive review of the mechanics of the
natural systems in primeval times. Despite his valiant
attempts to do this there are inherent defects. The
interrelationships of animals and plants were not well
understood during Shelford’s time and consequently
are poorly documented. In addition, the book fails to
present quantitative data on animal populations and
their food habits, and plant population dynamics.
The book is accented with maps, graphs, tables and
photographs. The references are extensive, and are
useful to those who wish to examine some of the
earlier works in ecology.
I recommend this book to biologists who are well
read in general ecology and who are capable of
recognizing its inadequacies.
PAUL A. GRAY
Hough, Stansbury, Michalski Ltd., 1265 Arthur Street,
Suite 409, Thunder Bay, Ontario P7E 6E7
groups in the areas of chemistry, plant physiology,
toxicology, genetics, and ecology with economics. A
group of 35 internationally recognized experts par-
ticipated in the symposium. Unlike so many scientific
symposia, this one was results-oriented. Summary
papers with recommendations were prepared by each
of the working groups. All of the background papers
and summaries are given, along with a short overall
conclusions and recommendations section.
The chemistry section and a considerable propor-
1979
tion of the toxicology section are devoted principally
to tetrachlorodibenzo-p-dioxin, the notorious TCDD.
TCDD 1s an incidental contaminant formed during
the manufacture of trichlorophenol, which in turnisa
precursor of the herbicides 2,4,5-T and 2,4,5-TP
(fenoprop). The bulletin gives an interesting account
of the toxicological problems (real or imagined) from
phenoxy acid (largely 2,4-D) use in Sweden. One of
the more controversial aspects was the suspected
poisoning of a reindeer herd in 1970. The veracity of
other alleged cases of poisoning of humans, wildlife,
fish, and bees are discussed in some detail. In the
section on ecology and economics there is a thorough
coverage of the effects of herbicide use in silviculture
on vegetation, but not much information is given on
how these effects relate to animal or insect ecology
(i.e., habitat changes), in spite of the title. The
importance of this subject is recognized, however, in
the summary and recommendations. A good review of
the effect of phenoxy herbicides on soil organisms 1s
given.
According to the National Research Council of
Canada document on the same subject (Phenoxy
MISCELLANEOUS
BOOK REVIEWS
461
Herbicides, their Effects on Environmental Quality,
NRCC Number 16075), several million kilograms of
2,4-D are used in Canada, largely for weed control in
the prairies. In the order of 50,000 kilograms (1973-
1974) of 2,4,5-T are used for brush control on
rights-of-way, mainly in the eastern provinces. This
bulletin therefore addresses a situation in Canada of
similar magnitude and import to that in Sweden.
Although there are gaps in coverage (in large part due
to a lack of information, I suspect), this volume is one
of the best overall sources of information available on
the subject. The many sections on TCDD have
relevance apart from phenoxy herbicides, because
trichlorophenol manufacture and dumping have
created more TCDD problems than 2,4,5-T use. The
summaries and recommendations are succinct and
well-balanced from an overall risk-benefit viewpoint.
R.J. NORSTROM
Wildlife Toxicology Division, Canadian Wildlife Service,
Ottawa, Ontario KIA 0E7
Last of the Naturalists: the career of C. Hart Merriam
By Keir B. Sterling. 1977. Arno Press, New York. 472 pp.,
illus. U.S. $23.00.
This book should be perused by those interested in
the formation and early activities of the American
Ornithologists Union, National Geographic Society,
American Society of Mammalogists, and U.S. Fish
and Wildlife Service, all of which Merriam
(1855-1942) was directly involved in founding.
Sterling has exhaustively researched the life of
Merriam using file letters to and from Merriam,
congressional documents where Merriam partici-
pated, journal and newspaper articles, interviews with
people who knew him, and other detailed records to
produce a fine biographical work that doesn’t stop
with information on Merriam but includes per-
sonalities of people that were around him.
This revised book, first issued in 1974, is part of the
Natural Sciences in America collection. The text is clut-
tered with quite a few typographical errors: entire
words and even sentences seem to be missing in some
spots. The chapter on speciation is poorly written. As
well, errant definitions occur, suchas Bergmann’s rule
(p. 205). The Elliott Coues —- Merriam vendettas are
probably: not fully covered. Perhaps a better title
would be The First American Wildlife Biologist or
Mammalogist, as the book’s present title implies that
the Leopolds and Muries were not naturalists.
Merriam’s career included being an amateur
naturalist, a medical school graduate, practicing
physician and professional biologist with the govern-
ment. He worked under the vigilance and influence of
Joel A. Allen, Spencer F. Baird, Jean B. Lamarck,
Charles V. Riley, and Theodore Roosevelt. He
influenced the professional lives of Vernon Bailey,
William Brewster, Frank M. Chapman, Albert K.
Fisher, Joseph Grinnell, Francis Harper, Henry W.
Henshaw, Waldo L. McAtee, Edward W. Nelson, and
Wilfred Osgood, many of whom worked for the U.S.
Biological Survey under his leadership.
Merriam was highly interested in the biogeography
and systematics of mammals and birds (he collected
thousands of specimens); published his first paper at
age 17, became an authority on birds at 22; was the
United States’ first full-time professional ornitholo-
gist; did not “follow the herd”; scorned religion; liked
to eat meat of cat, eagle, and skunk; had yearly field
experience throughout his life; believed that species
were basically fixed but all were not yet discovered:
named and described many new genera, species, and
subspecies, being a splitter when it came to zoological
462 THE CANADIAN FIELD-NATURALIST
nomenclature; put forth the life-zone theory, feeling
that temperature was the ultimate factor affecting the
distribution of plants and animals; worked intensively
with gophers and bears; thought life was too short for
memorizing terminology, believing that practical
application was pertinent to all work; felt that
microscopic work in the laboratory was emphasized
too much over field work in college; did not get along
with others whose drives were as great as his; was
involved in the still controversial killing of seals in the
north; could not receive criticism well; tried to get deer
farming initiated in areas where cows could not graze;
liked art and automobiles; despised the words biota
and ecology; was against conservation movements
that were not scientifically based; had a special ability
to obtain capable men to work for him at the
Le Naturaliste Canadien, Index
By G.-W. Corrivault and P. Morisset. 1979. Les Presses de
PUniversité Laval, Québec. 352 pp. $10.
Readers of The Canadian Field-Naturalist are
familiar with the problems of data retrieval from a
long series of volumes. The search for specific data
through so many annual indices Is not only tedious, it
may also be a poor use of one’s time. Because annual
indices are not equally thorough and comprehensive,
some being rather skimpy, one is never sure that
significant data has not escaped such a search.
Cumulative indices are the usual solution to these
problems, but not every journal has been able to
prepare and publish these.
Le Naturaliste Canadien is now in its 105th volume
and its data retrieval problems are quite similar to
those of The Canadian Field-Naturalist, except for
the existence of a series of cumulative indices of which
the present one is the fourth. A cumulative index
covering the first 20 volumes was published at the end
of volume 20. A second cumulation was printed at the
end of volume 54 and covers volumes 21-54. A
cumulated author index for volumes 1-81 will be
found at the beginning of volume 82.
The new cumulation carries the coverage forward to
volume 100. It is a separate publication, not a
particular issue of the serial. It is published in two
parts. The first part is analphabetized author index of
Vol. 93
Biological Survey; served office in many scientific
societies; Was most particular about the proper
preparation of skins and skulls; became interested in
Indians in the early 1900s, mostly ignoring or even
terming as useless most other mammals for the
remainder of his life; later in life turned against the
Biological Survey’s predator poisoning campaign
which he started; was never considered an intellectual;
and never accepted the developments that progressed
in genetics. He was the father of modern
Mammalogy.
RICHARD M. ZAMMUTO
Department of Zoology, University of Western Ontario,
London, Ontario N6A 5B7
2504 papers found in volumes | to 100. By design the
author index is not thorough: many minor items of
fleeting value have been omitted, such as notices from
the editor to his readers, some letters to the editor,
society notices, and the like.
The second part deals only with volumes 55 to 100.
It is a subject and taxon index with double references.
The bracketed numbers refer to the bibliography and
are followed by the usual volume and page references.
There are more than 60000 individual entries.
Botanists will be especially interested by this index.
There is very little botany in the first 50 volumes of Le
Naturaliste Canadien. But in the 1920s and 1930s
botanical papers gradually gained in numbers and
import. More than 8000 plant names occur in the
index, including nearly all the vascular plants of
Canada, Greenland, and Alaska; also a majority of
Canadian mosses and lichens, and a fair showing of
algae and fungi.
Taxonomic innovations are underlined in the
index: there are more than a thousand of them anda
good half of them are plant names.
B. BOIVIN
Biosystematics Research Institute, Agriculture Canada,
Ottawa, Ontario KIA 0C6
Se
1979
NEW TITLES
Zoology
The alligator: king of the wilderness. 1977. By William and
Ellen Hartley. Nelson, New York. 175 pp. US $7.95.
+ Animal behavior: an evolutionary approach. 1979. By John
Alcock. Second edition. Sinaver, Concord, Maine. xii +
532 pp., illus. US $16.
Animals of the oceans: the ecology of marine life. 1977. By
M. Angel and T. Harris. Two Continents, New York. 156
pp., illus. US $10.95.
Arachnology. 1978. Edited by P. Merrett. Proceedings of
the Symposia of the Zoological Society of London,
Number 42, Exeter, England, July 1977. Academic, New
York. xxxil + 530 pp., illus. US $46.50.
Eloquent animals: a study in animal communication. 1978.
By Flora Davis. Coward, McCann, and Geoghegan, New
York. 223 pp. US $8.95.
Field guide to seashells of the world. 1978. By Gert Lindner.
Van Nostrand Reinhold, New York. 271 pp., illus. Cloth
US $12.95; paper US $8.95.
Fifty common birds of Oklahoma and the south great
plains. 1977. By G.M. Sutton. University of Oklahoma
Press, Norman. xii + 113 pp. US $7.95.
Insect magic. 1978. By Kjell B. Sandved and Michael G.
Emsley. Viking, New York. 128 pp., illus. US $16.95.
t+Management and biology of Pacific flyway geese. 1979.
Edited by Robert L. Jarvis and James C. Bartonek. Oregon
State University Book Stores, Corvallis, Oregon. 346 pp.,
illus. Paper US $5.50.
Primates of South Asia: ecology, sociobiology and be-
havior. 1977. By M.L. Roonwal and S.M. Mohnot.
Harvard University Press, Cambridge, Massachusetts.
xx + 421 pp., illus. US $25.
Responses of Peary Caribou and Muskoxen to helicopter
harassment. 1979. By Frank L. Miller and Anne Gunn.
Occasional Paper Number 40, Canadian Wildlife Service,
Edmonton. 90 pp. Free.
Sea squirts of the Atlantic continental shelf from Maine to
Texas. 1979. By Harold H. Plough. Johns Hopkins
University Press, Baltimore. ix + 118 pp., illus. US $20.
t+Studies on Atlantic halocyprid ostracods. 1979. By Martin
V. Angel. Pergamon Press, Elmsford, New York. 128 pp..
illus. US $27.50.
The vertebrates: their forms and functions. 1978. By
Charles G. Crispens, Jr. Thomas, Springfield, Illinois.
x + 209 pp., illus. US $15.50.
BOOK REVIEWS
463
Water birds of California. 1977. By Howard L. Cogswell.
University of California Press, Berkeley. vi + 399 pp.., illus.
US $5.75.
*Wolf ecology and prey relationships on Isle Royale. 1978.
By Rolf Olin Peterson. National Park Service Scientific
Monograph Series Number 11, United States Department
of the Interior, Washington. 211 pp. Paper.
The world of worms. 1978. By Dorothy Hinshaw Patent.
Holiday House, New York. 11 + 124 pp., illus. US $6.95.
Botany
Botany: basic concepts in plant biology. 1978. By Terry L.
Hufford. Harper and Row, New York. xviii + 535 pp., illus.
US $16.95.
Essays in plant taxonomy. 1978. Edited by H.E. Street.
Academic Press, New York. 304 pp. US $24.50.
*A guide to the literature on the herbaceous vascular flora of
Ontario. 1978. By James L. Hodgins. Revised edition.
Botany Press, 90 Wolfrey Avenue, Toronto. 73 pp. Paper
$4.
*How to prepare common wild foods. 1978. By Darcy
Williamson. D. Williamson, Box 1032, McCall, Idaho
83638. 110 pp., illus. Paper US $6.95.
Introduction to the algae: structure and reproduction. 1978.
By Harold C. Bold and Michael J. Wynne. Prentice-Hall,
Englewood Cliffs, New Jersey. xiv + 706 pp., illus. US $24.
The physiology of woody plants. 1979. By Paul J. Kramer
and Theodore T. Kozlowski. Academic Press, New York.
Plant strategies and vegetation processes. 1979. By J.P.
Grime. Wiley-Interscience, New York. 264 pp. Cloth US
$26.50; paper US $10.95.
Plantcraft: a guide to the everyday use of wild plants. 1978.
By Richard Mabey. Illustrated by Marjorie Blamey.
Universe, New York. 176 pp., illus. US $10.
The powdery mildews. 1978. Edited by D.M. Spencer.
Academic Press, New York. 586 pp. US $54.75.
+The rare vascular plants of Saskatchewan. 1979. By Robert
V. Maher, George W. Argus, Vernon L. Harms, and John
H. Hudson. Syllogeus Number 20. National Museum of
Natural Science, Ottawa. 55 pp. + maps English + 55 pp.
French. Free.
River plants: the macrophytic vegetation of watercourses.
1978. By S.M. Haslam. Illustrated by P.A. Wolseley.
Cambridge University Press, New York. xii + 396 pp., illus.
Cloth US $62.50; paper US $14.95.
464
Vegetation and production ecology of an Alaska arctic
tundra. 1978. Edited by L.L. Tieszen. Springer-Verlag, New
York. xvill + 217 pp., illus. US $34.80.
Environment
Adaptive environmental assessment and management.
1978. Edited by C.S. Holling. IHASA International Series
on Applied Systems Analysis, 3. Wiley-Interscience, New
York. xx + 378 pp., illus. US $16.50.
* Aleta Karstad’s Canadian nature notebook. 1979. By Aleta
Karstad. McGraw-Hill Ryerson, Toronto. 144 pp., illus.
SIZ295:
Biology and ethics: reflections inspired by a UNESCO
symposium. 1978. By Bruno Ribes. UNESCO (Canadian
distributor Renouf Publishing, Montreal). 202 pp. $14.
Coral reefs: research methods. 1978. Edited by D.R.
Stoddart and R.E. Johannes. Monographs on Oceano-
graphic Methodology, 5. UNESCO (Canadian distributor
Renouf Publishing, Montreal). xv + 581 pp. $33.60.
The ecology of fossils: an illustrated guide. 1978. Edited by
W.S. McKerrow. MIT Press, Cambridge, Massachusetts.
384 pp., illus. US $22.50.
Environmental assessment: approaching maturity. 1978.
By Selina Bendix and Herbert R. Graham. Ann Arbor
Science, Ann Arbor, Michigan. xi + 288 pp. US $20.
The environmental impact statement process: a guide to
citizen action. 1978. By Neil Orloff. Information Resources
Press, Washington. 200 pp. US $10.95.
Exploring nature with your child. 1977. By Dorothy
Shuttlesworth. Abrams, New York. 240 pp., illus. US
$18.50.
Freshwater wetlands: ecological processes and manage-
ment potential. 1978. Edited by Ralph E. Good, Dennis F.
Whigham, and Robert L. Simpson. Academic Press, New
York. 392 pp. US $17.50.
Handbook of environmental data and ecological par-
ameters. 1979. Edited by S.E. Jorgensen. Pergamon Press,
Elmsford, New York. 1100 pp. US $150.
Handbook of environmental data on organic chemicals.
1977. By Karel Verschueren. Van Nostrand Reinhold, New
York. vit+ 659 pp. US $37.50.
An introduction to population ecology. 1978. By G. Evelyn
Hutchinson. Yale University Press, New Haven. xi + 260
pp. US $17.50.
Investigating ecology. 1978. By E.H. Blaustein and R.
Blaustein. Arco, New York. 140 pp. Cloth US $11.95: paper
US $7.95.
THE CANADIAN FIELD-NATURALIST
Vol. 93
Key works to the fauna and flora of the British Isles and
northwest Europe. 1978. Edited by G.J. Kenrich, D.L.
Hawksworth, and R. Sims. Academic Press, New York. 192
pp. US $16.15.
Marine biota of the NE Pacific: a bibliography emphasizing
systematics and distribution. 1978. By W.C. Austin and
M.M. Deutsch. Khoyatan Marine Laboratory, Cowichan
Station, British Columbia. Variously paged. Paper $16.
Oceanography: concepts and history. 1978. Edited by
Margaret B. Deacon. Benchmark Papers in Geology,
Volume 35. Dowden, Hutchinson, and Ross (Distributed
by Academic Press, New York). 416 pp. US $25.
The pollution of flowers by insects. 1978. Edited by A.J.
Richards. Academic Press, New York. 262 pp. US $26.
Sampling design and statistical methods for environmental
biologists. 1979. By Roger H. Green. Wiley-Interscience,
New York. 272 pp. US $19.95.
Tropical forest ecosystems: a state of knowledge report. By
UNESCO, UNEP, and FAO. Natural Resources Research,
XIV. UNESCO (Canadian distributor Renouf Publishing,
Montreal). 683 pp. Paper $53.50; cloth $66.50.
Miscellaneous
Suicide or survival? The challenge of the year 2000. 1978.
Various authors. UNESCO (Canadian distributor Renouf
Publishing, Montreal). 192 pp. $14.
Astronomy: from the earth to the universe. 1979. By J.M.
Pasachoff. Saunders, New York. 476 pp., illus. $15.95
(Canadian $19.15).
+ Decision making for national parks in Canada north of 60°.
1978. By Terry Fenge. Working Paper Number 3, Presi-
dent’s Committee on Northern Studies. University of
Waterloo, Waterloo. 11 + 58 pp. Paper, no price given.
+The development of tourism and its potential futures in
Canada north of 60° with implications for national parks
and related reserves. 1978. By Bryan Smale. Working Paper
Number 1, President’s Committee on Northern Studies.
University of Waterloo, Waterloo. 11 + 61 pp. Paper, no
price given.
+Elements of park and recreation administration. 1979. By
Charles E. Doell and Louis F. Twardzik. Fourth edition.
Burgess Publishing, Minneapolis. vi + 367 pp. US $13.95.
+An international comparison of policies and institutional
arrangements for national parks and related reserves in
hinterland areas. 1978. By Julia Gardner. Working Paper
Number 2, President’s Committee on Northern Studies.
University of Waterloo, Waterloo. 11 + 61 pp. Paper, no
price given.
*Assigned for review
+ Available for review
Index to Volume 93
Compiled by W. HARVEY BECK
Aalders, L.E., 415
Acer nigrum, 34
rubrum, 34
saccharum, 32
Achillea borealis, 184
Actitis macularia, 188
Adamcik, R.S., A. W. Todd, and L. B. Keith. Demographic
and dietary responses of Red-tailed Hawks during a
Snowshoe Hare fluctuation, 16
Agabus sp., 171
Aggresion in Tree Swallows, Influence of weather on, 437
Agonum decentis, 80
Agrohordeum macounii, 166
Agropyron violaceum, 355
yukonense, 184
Agrostis borealis, 355
exarata, 163
Alaska, 77
Alaska, Fall foods of Common Snipe on the Copper River
Delta, 171
Alaska, Response of wintering Moose to mechanical habitat
rehabilitation in, 19]
Alberta, 16, 67, 176, 272, 363, 411
Alberta, Black-necked Stilts nesting near Edmonton, 68
Alberta, Characteristics of Peregrine Falcons migrating
through central, 1969-1978, 296
Alberta, Habitat utilization and population densities of the
amphibians of northeastern, 144
Alberta, Largest Gray Wolf skulls found in, 308
Alberta, Life history characteristics of Little Brown Bats
(Myotis lucifugus) in, 243
Alberta, Nesting of Calliope Hummingbird in Kananaskis
Provincial Park, 449
Alberta, Probable hybrids of Cinnamon X Blue-winged Teal
from southern, 316
Alberta, Reproductive biology of the Big Brown Bat
(Eptesicus fuscus) in, 48
Alberta, Taxonomy, distribution, and ecology of the cliff-
brake ferns ( Pe//aea: Polypodiaceae) in, 288
Alberta, Winter denning of the Striped Skunk in, 252
Alberta, Winter feeding by Porcupines in montane forests of
southwestern, 405
Alces alces gigas, 191
Allium schoenoprasum var. sibiricum, 167
Amblystegium juratzkanum, 140
Ambystoma laterale, 194
maculatum, 194
Amelanchier alnifolia, 167, 184
arobrea, 32
Amphibians of northeastern Alberta, Habitat utilization and
population densities of the, 144
Anaphalis margaritacea var. subalpina, 163
Anas cyanoptera, 316
discors, 316
platyrhynchos, 55
Andrews, S., 449
Andropogon gerardi, 35
Anenome drummondtii, 184
multifida, 184
narcissiflora s. lat., 263
parviflora, 184
Anomodon minor, 140
Antennaria Friesiana ssp. compacta, 265
neoalaskana, 265
Aquila chrysaetos, 367
Arabis canadensis, 36
Aralia nudicaulis, 163
Arctic Canada, Ivory Gull colonies in southeastern Elles-
mere Island, 173
Arctostaphylos rubra, 184
uva-ursi, 184
Ardea herodias, 439
Arenaria interpres, 188
melanocephala, 188
Arnica alpina, 184
cordifolia, 169
Artemisia alaskana, 184
frigida, 184
hyperborea, 184
rupestris, 184
Article or note?, 337
Asclepias exaltata, 36
Asio flammeus, 411
Asplenium platyneuron, 36
Aster alpinus, 355
alpinus ssp. vierhapperi, 169, 265
ciliolatus, 163
ericoides, 35
macrophyllus, 33
modestus, 169
pilosus, 36
Sagittifolius, 33
sibiricus, 184
yukonense, 184
Astragalus americanus, 168
umbellatus, 184
Athyrium filix-femina ssp. cyclosorum, 165
Atrichum tenellum, 431
Avena sativa, 285
Avocet, American, 68
Award, FON Conservation, for The Canadian Field-
Naturalist, 452
Award, Robert Lister receives, 337
Bacidia chlorococca, 140
Badger, 365
Bailey, R.O. Wild Mallard stocking in a large marsh
habitat, 55
Ball, P.W. Thaspium trifoliatum (Meadow-parsnip) in
Canada, 306
Barclay-Estrup, P. and R. A. Sims. Epiphytes on White
Elm, Ul/mus americana, near Thunder Bay, Ontario,
139
465
466
Barkworth, M. E. reviews by, 104, 212
Bassett, I. J., reviews by, 102, 103
Bat, Big Brown, 245
Keen’s, 247
Bat, Big Brown, (Eptesicus fuscus) in Alberta, Reproductive
biology of, 48
Bats, Little Brown, (Myotis lucifugus) in Alberta, Life
history of, 243
Beam, B., review by, 95
Bear, Black, 423
Beechey, T. J., review by, 100
Beetles, carabid, Examination of overwintering, for asso-
ciated mites, 79
Bellolio-Trucco, G., 431
Berberis nervosa, 128
Bidens cernua, 163
Biological flora of Canada. 1. Vaccinium angustifolium
Ait., Sweet Lowbush Blueberry, The, 415
Biology — the unknown science? (editorial), 6
Bird banding, Inland, 202
Bird, D. M., review by, 205
Bjorge, R. R., 252
Black Bear confrontation, Blue Grouse brood hen, 200
Blackbird, Brewer’s breeding in the Northwest Territories,
76
Blarina brevicauda, 315, 320
Blue Grouse brood hen — Black Bear confrontation, 200
Blueberry, Sweet Lowbush, The biological flora of Canada.
1. Vaccinium angustifolium Ait., 415
Bluebird society, 202
Bobbette, R. S. W., review by, 344
Boissonneau, A. N., review by, 213
Boivin, B., review by, 462
Bonasa umbellus, 18
Bosmina sp., 267
Botrychium virginianum var. europaeum, 163
Boxall, P. C., 411
Boxall, P. C. Interaction between a Long-tailed Weasel and
a Snowy Owl, 67
Boyd, H., review by, 105
Brachythecium relfexum, 140
salebrosum, 140
Brasenia schreberi, 389
Breeding areas and overnight roosting locations in the
northern range of the Monarch Butterfly (Danaus
plexippus plexippus) with a summary of associated
migratory routes, 41
Breeding in the Northwest Territories, Brewer’s Blackbird,
76
Brewster, G. J., 132
British Columbia, 70, 71, 200, 433
British Columbia, Giant Cow Parsnip (Heracleum mante-
gazzianum) on Vancouver Island, 82
British Columbia, Nesting of Horned Puffins in, 84
British Columbia, Virgin Douglas Fir forest on Saturna
Island, 126
Britton, D. M., review by, 345
Bromus ciliatus, 166
Brunton, D. F., review by, 347
Brunton, D. F., Taxonomy, distribution, and ecology of the
cliff-brake ferns (Pellaea: Polypodiaceae) in Alberta,
288
THE CANADIAN FIELD-NATURALIST
Vol. 93
Brunton, D. F., S. Andrews, and D. G. Paton. Nesting of
the Calliope Hummingbird in Kananaskis Provincial
Park, Alberta, 449
Buffalo, Black, (Osteichthyes: Catostomidae), First Can-
adian record of the, 304
Bufo hemiophrys, 144
Busby, D. G., 446
Buteo jamaicensis, 16, 367
swainsoni, 367
Butterfly, Monarch, (Danaus plexippus plexippus), Breed-
ing areas and overnight roosting locations in the
northern range of the, with a summary of associated
migratory routes, 41
By-laws of The Ottawa Field-Naturalists’ Club, Notice of
change to the, 337
Calamagrostis canadensis ssp. Lansdorffii, 262
deschampsiodes, 355
purpurascens, 184, 262
Calathus ingratus, 80
Calidris alpina, 188
bairdi, 188
mauri, 188
minutilla, 188
pusilla, 188
Calla palustris, 262, 389
Callitriche heterophylla, 389
Calosoma frigidum, 80
Cameron, R. D., K. R. Whitten, W. T. Smith, and D. D.
Roby. Caribou distribution and group composition
associated with construction of the Trans-Alaska Pipe-
line, 155
Campbell, C., 28
Campbell, R. W., H. R. Carter, and S. G. Sealy. Nesting of
Horned Puffins in British Columbia, 84
Canada, Man’s influence on potential nesting sites and
populations of swallows in, 371
Canada, Thaspium trifoliatum (Meadow-parsnip) in, 306
Canada, The biological flora of, |. Vaccinium angustifolium
Ait., Sweet Lowbush Blueberry, 415
Canada, The Oriskany sandstone outcrop and associated
natural features, a unique occurrence in, 28
Canada, Utricularia geminiscapa at Mer Bleue and range
extensions in eastern, 391
Canadian Field-Naturalist, FON Conservation Award for
The, 452
Candelaria concolor, 140
Canis latrans, 367
lupus, 308
l. irremotus, 308
l. occidentalis, 308
|. pambasileus, 308
Capella gallinago delicata, 171
Cardamine umbellata, 263
Carex aquatilis, 184
atherodes, 167
bigelowii, 355
canescens var. subloliacea, 262
concinna, 167
filifolia, 183
glacialis, 262
lapponica, 262
laxiflora var. gracillima, 36
1979
membranacea, 262
pedunculata, 34
pensylvanica, 33
Williamsii, 262
Caribou, Barren-ground, 155
Woodland, 71
Caribou distribution and group composition associated with
construction of the Trans-Alaska Pipeline, 155
Caribou, Distribution and movements of Selkirk, 1972-
1974, 71
Carpinus caroliniana, 34
Carter, H. R., 84
Carya cordiformis, 34
ovata, 32
Cassiope tetragona, 184
Castilleja caudata, 168
miniata, 163
Catfish, Flathead, First Canadian record of a, 179
Catling, P. M., 399
Catoptrophorus semipalmatus, 188
Catostomus commersoni, 267
Cavers, P. B., 378
Cedar, Eastern White, Thuja occidentalis, in western Nova
Scotia, Status of, 326
Celtis occidentalis, 34
Cerastium maximum, 261
Ceratophyllum demersum, 389
Cervus elaphus, 282
Cetraria ciliaris, 140
Chamaedaphne calyculata, 264
Chaoborus albipes, 267
Charadrius alexandrinus, 188
semipalmatus, 188
vociferus, 188
wilsonia, 188
Chipmunk, 320
Least, 314
Chlidonias niger, 276
Cicuta maculata var. angustifolia, 168
Cinclidium latifolium, 431
Cinna latifolia, 163
Circaea alpina, 163
Cirsium arvense, 284
Cladonia spp., 286
Clethrionomys gapperi, 314, 320
Cody, W. J., 163, 259
Cody, W. J., reviews by, 211, 212, 457
Collomia linearis, 168
Conopholis americana, 36
Constitution of the Ottawa Field-Naturalists’ Club, Notice
of motion to amend the, 338
Cook, F. R., 321
Cook, F. R. and J. Preston. Two-lined Salamander, Eurycea
bislineata, in Labrador, 178
Cormorant, Great, Wind-caused death of, 175
Cornus racemosa, 33
stolonifera, 284
Coyote, 367
Crappie, Black, in the Ottawa River, Seasonal growth,
food, and feeding habits of young-of-the-year, 232
Crataegus chrysocarpa, 34.
sp., 34
INDEX TO VOLUME 93
467
Crepis nana, 184
Croskery, P., reviews by, 93, 97, 107, 339
Crossman, E. J. and J. H. Leach. First Canadian record ofa
Flathead Catfish, 179
Crossman, E. J. and S. J. Nepszy. First Canadian record of
the Black Buffalo (Osteichthyes: Catostomidae), 304
Cutright, N. J., reviews by, 99, 207, 342, 456
Cyclops sp., 267
Cygnus olor, 433
Cypress Hills population of White-crowned Sparrows, Song
pattern of the, 272
Cypripedium calceolus var. planipetalum, 355
Cyr, A., review by, 106
Cystopteris fragilis ssp. fragilis, 261
Dagg, A. I., reviews by, 109, 217
Dale, H. M., 386
Danaus plexippus plexippus, Breeding areas and overnight
roosting locations in the northern range of the Monarch
butterfly with a summary of associated migratory
routes, 41
Dance, K. W., 180
Danthonia intermedia, 166
spicata, 34
Daphnia longispina, 267
David, N. Barn Owls in Quebec, 323
Davies, R.W. Dispersion of freshwater leeches (Hir-
udinoidea) to Anticosti Island, Quebec, 310
Dawe, N. K. and E. R. White. Giant Cow Parsnip ( Hera-
cleum mantegazzianum) on Vancouver Island, British
Columbia, 82
DDT, Eggshell thickness in American shorebirds before and
since, 187
Deer, Black-tailed, 130
Dekker, D. Characteristics of Peregrine Falcons migrating
through central Alberta, 1969-1978, 296
Deckker, D., R. Lister, T. W. Thormin, D. V. Weseloh, and
L. M. Weseloh. Black-necked Stilts nesting near Ed-
monton, Alberta, 68
Dendragapus obscurus, 200
Denning of the Striped Skunk in Alberta, Winter, 252
Densmore, R., 77
DesGranges, J.-L. Abandoned windmill used as a nesting
site by Great Blue Herons, 439
DesGranges, J.-L., review by, 94
Desmodium canadense, 34
dillenti, 35
paniculatum, 35
rotundifolia, 35
Desmognathus fuscus fuscus, 193
Development of young in Ontario Black Terns, Nesting
biology and, 276
Diaptomus oregonensis, 267
Dicranum fuscescens, 129
montanum, 140
Diets of Elk in Saskatchewan, Summer, autumn, and winter,
282
Dispersion of freshwater leeches (Hirudinoidea) to Anti-
costi Island, Quebec, 310
Disporum lanuginosum, 36
Distribution and group composition associated with con-
struction of the Trans-Alaska Pipeline, Caribou, 155
468
Distribution and habitats of four annual smartweeds in
Ontario, 378
Distribution and movement of Selkirk Caribou,
1974, 71
Distribution, Taxonomy, and ecology of the cliff-brake
ferns ( Pellaea: Polypodiaceae) in Alberta, 288
District of Mackenzie, Range extensions of vascular plants
in northern Yukon Territory and northwestern, 259
Draba alpina, 263
borealis, 263
crassifolia, 263
incerta, 263
Drepanocladus revolvens var. intermedius, 431
Drosera rotundifolia, 263
Dryas alaskensis, 263
drummondii, 167, 184
integrifolia, 184
octopetala ssp. alaskensis, 263
Dryopteris fragrans, 262
spinulosa, 163
Dunn, E. H. Nesting biology and development of young in
Ontario Black Terns, 276
Eagle, Bald, 367
Golden, 367
Eagles, Bald, in central
summering, 443
Eagles, P. F. J., reviews by, 102, 347
Earthworm cocoons as a drift component in a southern
Ontario stream, 180
Ecology of the cliff-brake ferms (Pellaea: Polypodiaceae)
in Alberta, Taxonomy, distribution, and, 288
Editor’s report for 1978, 201
Edwards Y. Biology — the unknown science? (editorial),
6
Edwards, Y. Is biology unknown? (letter), 330
Eggshell thickness in American shorebirds before and since
DDT, 187
Egret, Snowy, in the Northwest Territories, 329
Egretta thula, 329
Elaphe obsoleta obsoleta, 28, 322
vulpina gloydi, 399
Elatine minima, 388
Eleocharis acicularis, 388
robbinsii, 389
Elk in Saskatchewan, Summer, autumn, and winter diets
of, 282
Ellesmere Island, Arctic Canada, Ivory Gull colonies in
southeastern, 173
Ellesmere Island, Ring counts in Salix arctica from, 81
Elm, White, Ulmus americana, near Thunder Bay, Ontario,
Epiphytes on, 139
Elodea canadensis, 389
Elymus glaucus, 129
macouniil, 166
Epilobium angustifolium, 184, 284
latifolium, 184
Epiphytes on White Elm, U/mus americana, near Thunder
Bay, Ontario, 139
Eptesicus fuscus, 245
f. pallidus, 48
Eptesicus fuscus, Reproductive biology of the Big Brown
Bat in Alberta, 48
1972-
New Brunswick, Decline of
THE CANADIAN FIELD-NATURALIST
Vol. 93
Equisetum hyemale var. affine, 166
spp., 284
Erethizon dorsatum, 405
Erigeron caespitosus, 184
compositus, 184
Philadelphicus, 169
Eriocaulon septangulare, 388
Eriophorum angustifolium ssp. triste, 262
russeolum var. albidum, 262
triste, 262
Eritrichium splendens, 264
Erskine, A. J. Man’s influence on potential nesting sites
and populations of swallows in Canada, 371
Etheostoma sp., 268
Eumeces fasciatus, Five-linked Skink, in Ontario, Eastern
limit of the, 321
Euphagus cyanocephalus, 76
Eurhynchium oreganum, 126
Eurycea bislineata, Ywo-lined Salamander in Labrador, 178
Eurycea bislineata bislineata, 194
Eutamias minimus, 314
Evernia mesomorpha, 140
Fagus grandifolia, 34
Fahselt, D., P. Maycock, G. Winder, and C. Campbell.
The Oriskany sandstone outcrop and associated natural
features, a unique occurrence in Canada, 28
Falco peregrinus, 296
p. anatum, 297
p. tundrius, 297
sparverius, 198
Falcons, Peregrine, migrating through central Alberta,
1969-1978, Characteristics of, 296
Feeding, Aquatic, by a Woodchuck, 309
Feeding habits of young-of-the-year Black Crappie in the
Ottawa River, Seasonal growth, food, and, 232
Feeding, Winter, by Porcupines in montane forests of
southwestern Alberta, 405
Festuca occidentalis, 129
ovina ssp. alaskana, 262
subulata, 129
Fetterolf, P. M. Common Garter Snake predation on Ring-
billed Gull chicks, 317
Fetterolf, P. M. Intraspecific food theft by the American
Kestrel, 198
Field research investigators, Proposals invited from, 201
Fir, Douglas, forest on Saturna Island, British Columbia,
Virgin, 126
Fissidens obtusifolius, 431
FON Conservation Award for The Canadian Field-Nat-
uralist, 452
Fontinalis sp., 388
Food and feeding of the Rainbow Smelt (Osmerus mordax)
in Lake Simcoe, Ontario, 266
Foods of Common Snipe on the Copper River Delta,
Alaska, Fall, 171
Food-storage behavior of Peromyscus maniculatus gracilis
and p. leucopus noveboracensis, Nesting and, 239
Forest, oak-hickory, 28
Fox, Red, 423
Foxall, R. A. One hundred years in perspective — the chang-
ing roles and objectives of The Ottawa Field-Natural-
ists’ Club, |
ITY .
Fragaria vesca, 129
Fraser, D. Aquatic feeding by a Woodchuck, 309
Fratercula corniculata, 84
Fraxinus americana, 32
Freddy, D.J. Distribution and movements of Selkirk
Caribou, 1972-1974, 71
Freedman, B. and P.M. Catling. Movements of sym-
patric species of snakes at Amherstburg, Ontario, 399
Freitag, R., 79
Frisch, T. and W. C. Morgan. Ivory Gull colonies in south-
eastern Eilesmere Island, Arctic Canada, 173
Frog, Boreal Chorus, 144
Wood, 144
Frullania bolanderi, 140
eboracensis, 140
Galium aparine, 33, 129
boreale, 184
Brandegei, 261
circaezans, 33
pilosum, 35
triflorum, 169
Gallinago paraguaiae, 189
Gammarus, 267
Garter Snake, Common, predation on Ring-billed Gull
chicks, 317
Gaultheria shallon, 126
Geranium maculatum, 33
richardsonii, 168
Geum aleppicum vat. strictum, 168
Glaucomys sabrinus, 320
Glossiphonia complanata, 310
Glyceria borealis, 388
Striata var. stricta, 166
Goodyera repens, 167
Goossen, J. P. and D. G. Busby. Occurrences of the Red
Phalarope in the prairie provinces and adjacent states,
446
Gordon, D. M. New localities for the Northern Spring
Salamander and the Four-toed Salamander in south-
western Quebec, 193
Goski, B. C., 259
Gray, P. A., reviews by, 104, 108, 215, 339, 460
Green, R. H., reviews by, 456
Grosbeak, Evening, Xanthochroism in the, 66
Grosbeaks, Pine, Interspecific vocal mimicry by, 436
Grouse, Ruffed, 17
Sharp-tailed, 17
Gruchy, C., review by, 206
Gull, Ivory, Colonies in southeastern Ellesmere Island,
Arctic Canada, 173
Gull, Lesser Black-backed, Larus fuscus, in Labrador
waters, 445
Gull, Ring-billed, chicks, Common Garter Snake predation
on, 317
Gulls, Herring, and Common Terns in northeastern Sas-
katchewan, Colonial nesting, 132
Gunson, J. R., 48, 243
Gunson, J. R. and R. M. Nowak. Largest Gray Wolf skulls
found in Alberta, 308
Gunson, J.R. and R.R. Bjorge. Winter denning of the
Striped Skunk in Alberta, 252
Gyrinophilus porphyriticus porphyriticus, 193
INDEX TO VOLUME 93
469
Haber, E. Utricularia geminiscapa at Mer Bleue and range
extensions in eastern Canada, 391
Habitat rehabilitation in Alaska, Response of wintering
Moose to mechanical, 191
Habitat selection by wintering Snowy Owls (Nyctea scan-
diaca), 176
Habitat use in Montana, Lynx movements and, 441
Habitat utilization and population densities of the amphibi-
ans of northeastern Alberta, 144
Habitats of four annual smartweeds in Ontario, Distri-
bution and, 378
Haematopus ostralegus bachmani, 188
oO. palliatus, 187 :
Haliaeetus leucocephalus, 367, 443
Hall, I. V., L. E. Aalders, N. L. Nickerson, and S. P. Vander
Kloet. The biological flora of Canada. 1.
Vaccinium angustifolia Ait., Sweet Lowbush Blueberry,
415
Hanson, J. M. and S. U. Qadri. Seasonal growth, food, and
feeding habits of young-of-the-year Black Crappie in
the Ottawa River, 232
Harder es D243
Harder, L. D. Winter feeding by Porcupines in montane
forests of southwestern Alberta, 405
Hare, Snowshoe, 19, 441
Harms, V. L., review by, 213
Harris, R. N. Influence of weather on aggresion in Tree
Swallows, 437
Hash, H.S., 441
Hawk, Red-tailed, 367
Swainson’s, 367
Hawks, Red-tailed, Demographic and dietary responses of,
during a Snowshoe Hare fluctuation, 16
Hedysarum alpinum, 184
mackenzii, 184
Helleiner, C. W. Xanthochroism in the Evening Grosbeak,
66
Helobdella stagnalis, 310
triserialis, 310
Hemidactylium scutatum, 193
Heracleum mantegazzianum, Giant Cow Parsnip on Van-
couver Island, British Columbia, 82
Herons, Great Blue, Abandoned windmill used as nesting
site, 439
Hesperiphona vespertina, 66
Hexagenia sp., 267
Hieracium scabriusculum, 169
Hierochloe alpina, 355
pauciflora, 355
Himanotopus mexicanus, 68, 188
Hippurus vulgaris, 309, 389
Hirudinoidea, Dispersion of freshwater leeches, to Anticosti
Island, Quebec, 310
Hirundo rustica, 374
Hoefs, M. Flowering plant phenology at Sheep Mountain,
southwest Yukon Territory, 183
Hogan, G. G., 175
Hordeum jubatum, 184
vulgare, 285
Hornocker, M. G., 441
Houston, C. S., review by, 207
470
Hummingbird, Calliope, in Kananaskis Provincial Park,
Alberta, Nesting of, 449
Hummingbird, Rufous, 450
Humphreys, G., 63
Hunt, H. M. Summer, autumn, and winter diets of Elk in
Saskatchewan, 282
Hybrids of Cinnamon X Blue-winged Teal from southern
Alberta, Probable, 316
Hylocomium splendens, 126
Hypericum perforatum, 34
Hypnum circinale, 129
hamulosum, 431
Tchthyomyzon fossor, First record of the Northern Brook
Lamprey, in Nelson River drainage, Manitoba, 199
Ictiobus niger, 304
Interaction between a Long-tailed Weasel and a Snowy Owl,
67
Interactions between Snowy and Short-eared Owls in winter,
41]
Ireland, R. R. and G. B. Bellolio-Trucco. Mosses new to
Ontario and Quebec, 431
Tridoprocne bicolor, 372, 437
Is biology unknown? (letters), 330
Tsoetes macrospora, 388
Tsothecium stoloniferum, 129
IUCN, Books available from, 87
IUCN views on whale management, 88
Jaagumagi, R., review by, 209
Jacana spinosa, 188
Jenkins, R. E., 63
Juglans nigra, 34
Juncus arcticus spp. alaskanus, 262
arcticus spp. ater, sensu, 167
balticus var. alaskanus, 262
balticus var. littoralis, 167
militaris, 389
pelocarpus, 388
Juniperus virginiana, 34
Jyrkkanen, J. and D.G. Wright. First record of the
Northern Brook Lamprey, /chthyomyzon fossor, in the
Nelson River drainage, Manitoba, 199
eno, 1, 1.5 NG
Kennedy, A. J., review by, 218
Kestrel, American, Intraspecific food theft by the, 198
Katies be 187
King, L. L., review by, 344
Kirkland, C. J., 195
Kirkland, G.L., Jr. and C.M. Knipe. The Rock Vole
(Microtus chrotorrhinus) as a Transition zone species,
319
Kirkland, G. L., Jr., D. F. Schmidt, and C. J. Kirkland.
First record of the Long-tailed Shrew (Sorex dispar) in
New Brunswick, 195
Klein, H. G., 239
Knipe, C. M., 319
Kobresia myosuroides, 355
simpliciuscula, 355
Koehler, G.M., M. N. Hornocker, and H. S. Hash. Lynx
movements and habitat use in Montana, 441
Kott, E.. R. E. Jenkins, and G. Humphreys. Recent collec-
tions of the Black Redhorse, Moxostoma duquesnei,
from Ontario, 63
THE CANADIAN FIELD-NATURALIST
Vol. 93
Labrador, Two-lined Salamander, Eurycea bislineata, in,
178
Labrador waters, Lesser Black-backed Gull, Larus fuscus,
in, 445
Lamprey, Northern Brook, Jchthyomyzon fossor, in the
Nelson River drainage, Manitoba, First record of the,
199
Laporte, P., review by, 92
Lappula myosotis, 184
Larus argentatus, 132
delawarensis, 317
fuscus graellsii, 446
Larus fuscus, Lesser Black-backed Gull,
waters, 445
Lathyrus ochroleucus, 284
venosus, 284
each erates |79
Lecanora conizaea, 140
impudens, 140
Ledum decumbens, 355
groenlandicum, 264
palustre, 184
palustre ssp. groenlandicum, 264
Leeches, freshwater (Hirudinoidea), Dispersion of, to Anti-
costi Island, Quebec, 310
Lein, M.R. Song pattern of the Cypress Hills popula-
tion of White-crowned Sparrows, 272
Lein, M. R. and G. A. Webber. Habitat selection by winter-
ing Snowy Owls (Nyctea scandiaca), 176
Lein, M.R. and P. C. Boxall. Interactions between Snowy
and Short-eared Owls in winter, 411
Lemna trisulca, 262
Lepidozia reptans, 129
Lepraria membranacea, 140
neglecta, 140
Leptodora kindtii, 267
Lepus americanus, 16, 441
Lespedeza capitata, 35
intermedia, 35
Lewin, V., 144
Lien, J., 303
Life history characteristics of Little Brown Bats (Myotis
lucifugus) in Alberta, 243
Limosa fedoa, 188
Lindsay, J. E., review by, 458
Limnaea borealis, 184
borealis ssp. americana, 264
Linum perenne, 184
virginiana, 35
Lister, R., 68
Lister, Robert, receives award, 337
Listera cordata, 129
Lloyd, Hoyes, 1888-1978, Tribute to, 331
Lobaria pulmonaria, 140
Lobelia dortmanna, 388
Lophocolea bidentata, 129
Lunda cirrhata, 84
Lupinus arcticus, 184
Luzula arctica, 262
arcuata s. lat., 262
confusa, 355
multiflora s. lat., 262
in Labrador
IQ)
nivalis, 262
parviflora, 262
Wahlenbergii ssp. Wahlenbergii, 261
Lycopodium annotinum ssp. pungens, 261
complanatum, 261
Lynx canadensis, 441
Lynx movements and habitat use in Montana, 441
MacCrimmon, H. R. and R. W. Pugsley. Food and feeding
of the Rainbow Smelt (Osmerus mordax) in Lake
Simcoe, Ontario, 266
Mackenzie, District of, Range extensions of vascular plants
in northern Yukon Territory and northwestern, 259
Mahon, R., review by, 349
Maianthemum canadense, 33
Mallard, Wild, stocking in a large marsh habitat, 55
Manitoba, 55, 75, 446
Manitoba, First record of the Northern Brook Lamprey,
Ichthyomyzon fossor, in the Nelson River drainage,
199
Marmota monax, 309
Martell, A. M. Relative efficiencies of Museum Special,
Victor, and Holdfast traps for sampling small mammal
populations, 313
Martin, K. Common Garter Snake predation on Robin
nestlings, 70
Martin, Purple, 371
Matteuccia struthiopteris var. pensylvanica, 163
Maycock, p., 28
McAllister, D. E., review by, 208
McDonald, H. G., review by, 454
McKeating, G. B., review by, 210
McKelvey, R. W. Swans wintering on Vancouver Island,
1977-1978, 433
MecNicholl, M. K. Communal roosting of Song Sparrows
under snowbank, 325
MeNicholl, M. K., reviews by, 98, 455
McNicholl, M. K., Robert Lister receives award, 337
MeNicholl, M. K. and G. G. Hogan. Wind-caused death of
Great Cormorant, 175
Meadow-parsnip, Thaspium trifoliatum, in Canada, 306
Medicago sativa, 285
Meeting, Annual Business, Notice of The Ottawa Field-
Naturalists’ Club, 452
Melospiza melodia, 325
Mephitis mephitis, 252
Merdsoy, B., J. Lien, and A. Storey. Extralimital record of
a Narwhal (Monodon monoceros) in Hall’s Bay, New-
foundland, 303
Mertensia paniculata, 184
Michener, G. R. Yearly variations in the population dynam-
ics of Richardson’s Ground Squirrels, 363
Mickelson, P. G., 171
Micropalama himantopus, 188
Microsorex hoyi, 315
Microtus chrotorrhinus, 314
pennsylvanicus, 18, 315, 320, 412
Microtus chrotorrhinus, The Rock Vole, as a Transition
zone species, 319
Miller, G.E. and H.M. Dale. Apparent differences in
aquatic macrophyte floras of eight lakes in Muskoka
District, Ontario from’ 1953 to 1977, 386
Mimicry by Pine Grosbeaks, Interspecific vocal, 436
INDEX TO VOLUME 93 471
Mimulus guttatus, 168
Mink, 276
Minuartia dawsonensis, 167
groenlandica, 355
Mites, Examination of overwintering carabid beetles for
associated, 79
Mnium cuspidatum, 140
Monarda fistulosa, 35
Moneses uniflora, 264
Monodon monoceros, Narwhal, in Hall’s Bay, Newfound-
land,Extralimital record of a, 303
Montana, Lynx movements and habitat use in, 44]
Moore, M. I., review by, 346
Mooreobdella fervida, 310
Moose, Response of wintering, to mechanical habitat
rehabilitation in Alaska, 191
Morgan, W. C., 173
Morrison, M. L. and L. F. Kiff. Eggshell thickness in Ameri-
can shorebirds before and since DDT, 187
Mosses new to Ontario and Quebec, 431
Mouse, Deer, 239, 314, 320, 412
Meadow Jumping, 314, 320
White-footed, 239
Woodland Jumping, 239, 320
Movements and habitat use in Montana, Lynx, 441
Movements of Selkirk Caribou, 1972-1974, Distribution
and, 71
Movements of sympatric species of snakes at Amherst-
burg, Ontario, 399
Moxostoma duquesnei, Recent collections of the Black
Redhorse from Ontario, 63
Moxostoma erythrurum, 63
macrolepidotum, 64
Muhlenbergia mexicana, 163
Munro, D.A., Tribute to Hoyes Lloyd, 1888-1978, 331
Murphy, D., review by, 453
Muskrat, 276
Muskrat in southeastern New Brunswick, Unusually late
pregnancy of a, 440
Mustela frenata, 67, 365, 412
nivalis, 68
vison, 276
Myosotis alpestris, 184
Myotis keenti, 247
lucifugus lucifugus, 243
Myotis lucifugus, Little Brown Bats, Life history char-
acteristics, in Alberta, 243
Myriophyllum farwelli, 389
tenellum, 389
Myurella tenerrima, 431
Nagy, J. A., A. M. Pearson, B. C. Goski, and W. J. Cody.
Range extensions of vascular plants in northern Yukon
Territory and northwestern District of Mackenzie,
259
Najas flexilis, 388
Napaeozapus insignis, 315, 320
Narwhal (Monodon monoceros) in Hali’s Bay, Newfound-
land, Extralimital record of a, 303
Nepszy, S. J., 304
Nesting and food-storage behavior of Peromyscus mani-
culatus gracilis and P. leucopus noveboracensis, 239
472
Nesting, biology and development of young in Ontario Black
Terns, 276
Nesting of Horned Puffins in British Columbia, 84
Nesting of the Calliope Hummingbird in Kananaskis Pro-
vincial Park, Alberta, 449
Nesting site, Abandoned windmill used as a, by Great
Blue Herons, 439
Nesting sites, potential, and populations of swallows in
Canada, Man’s influence on, 371
Nettleship, D. N., review by, 95
New Brunswick, 437
New Brunswick, Decline of summering Bald Eagles in
central, 443
New Brunswick, First record of the Long-tailed Shrew in,
195
New Brunswick, Unusually late pregnancy of a Muskrat in
southeastern, 440
New York, 239, 319
Newfoundland, Extralimital record of a Narwhal (Monodon
monoceros) in Hall’s Bay, 303
Newt, Red-spotted, 195
Nickerson, N. L., 415
Norstrom, R. J., review by, 460
Northcott, T. H., review by, 342
Northwest Territories, 81
Northwest Territories, Brewer’s Blackbird breeding in the,
76
Northwest Territories, Snowy Egret in the, 329
Notophthalmus viridescens, 195
Notropis atherinoides, 266
Nova Scotia, 67
Nova Scotia, Status of Eastern White Cedar,
occidentalis, in western, 326
Nowak, R. M., 308
Numenius americanus, 188
phaeopus, 188
Nuphar variegatum, 309, 389
Nyctea scandiaca, 67, 411
Nyctea scandiaca, Snowy Owls, Habitat selection by winter-
ing, 176
Nycticryphes semicollaris, 189
Nymphaea ordorata, 389
Nymphoides cordatum, 389
Odocoileus hemionus columbianus, 130
Olor buccinator, 433
Olynyk, J. E.and R. Freitag. Examination of overwintering
adult carabid beetles for associated mites, 79
Ondatra zibethicus, 276
z. zibethicus, 440
Ontario, 28, 79, 198, 306, 309
Ontario and Quebec, Mosses new to, 431
Ontario Black Terns, Nesting biology and development of
young in, 276
Ontario, Distribution and habitats of four annual smart-
weeds in, 378
Ontario, Eastern limit of the Five-lined Skink, Eumeces
fasciatus, in, 321
Ontario, Epiphytes on White Elm, U/mus americana, near
Thunder Bay, 139
Ontario, Food and feeding of the Rainbow Smelt (Osmerus
mordax) in Lake Simcoe, 266
Thuja
THE CANADIAN FIELD-NATURALIST
Vol. 93
Ontario from 1953 to 1977, Apparent differences in aquatic
macrophyte floras of eight lakes in Muskoka District,
386
Ontario, Movements of sympatric species of snakes at
Amherstburg, 399
Ontario, Recent collections of the Black Redhorse, Moxo-
stoma duquesnei, from, 63
Ontario, Some new and interesting vascular plant records
from northern, 355
Ontario stream, Earthworm cocoons as a drift component in
a southern, 180
Oriskany sandstone outcrop and associated natural features,
a unique occurrence in Canada, The, 28
Ornithologists, Societies and amateur, 88
Orthothecium strictum, 431
Orthotrichum obtusifolium, 140
speciosum, 140
strangulatum fo. lescurii, 432
Oryzopsis asperifolia, 163
Osmerus mordax in Lake Simcoe, Ontario, Food and feed-
ing of the Rainbow Smelt, 266
Osmorhiza depauperata, 163
obtusa, 168
Ostrya virginiana, 34
Ottawa Field-Naturalists’ Club Annual Business Meeting of
The, 452
Ottawa Field-Naturalists’ Club
Auditor’s report, 226
Balance sheet, 227
Minutes of the ninety-ninth annual business meeting,
DDB}
Report of council, 224
Statement of income and expenditure, C.F.N., 229
Statement of income and expenditure, O.F.N.C., 228
Ottawa Field-Naturalists’ Club, Call for nominations for
the council of The, 353
Ottawa Field-Naturalists’ Club. Notice of change to the by-
laws of The, 337
Ottawa Field-Naturalists’ Club, Notice of motion to amend
the constitution of The, 338
Ottawa Field-Naturalists’ Club, One hundred years in
perspective — the changing roles and objectives of, The,
|
Ottawa River, Seasonal growth, food, and feeding habits of
young-of-the-year Black Crappie in the, 232
Ouellet, H., reviews by, 210, 340
Owl, Snowy, Interaction between a Long-tailed Weasel and
a, 67
Owls, Barn, in Quebec, 323
Owls, Snowy and Short-eared, in winter, Interactions
between, 411
Owls, Snowy, (Nyctea scandiaca), Habitat selection by
wintering, 176
Oxytropis borealis, 264
deflexa var. foliolosa, 168
deflexa var. sericea, 264
glutinosa, 264
huddelsonii, 184
nigrescens ssp. pygmaea, 264
pygmaea, 264
viscida, 184
NTS
Oystercatcher, “American”, 189
“black”, 189
Pagophila eburnea, 173
Paleoclimate, Natural fires as an index of, 116
Panicum lanuginosum var. praecocius, 36
linearifolium, 35
Papaver Hultenii, 263
Keelei, 263
Macounii, 263
Parietaria pensylvanica, 163
Parker, G. R. Unusually late pregnancy of a Muskrat in
southeastern New Brunswick, 440
Parmelia saxatilis, 140
sulcata, 140
Parsnip, Giant Cow, ( Heraclium mantegazzianum) on Van-
couver Island, British Columbia, 82
Partridge, Gray, 412
Paton, D. G., 449
Pearson, A. M., 259
Pedicularis flammea, 361
labradorica, 361
lapponica, 264, 361
spp., 186
sudetica, 361
Pedioecetes phasianellus, 18, 74
Pellaea atropurpurea, 288
occidentalis, 288
suksdorfiana, 288
Pellaea (Polypodiaceae) in Alberta, Taxonomy, distribu-
tion, and ecology of the cliff-brake ferns, 288
Pentaneuris monilis, 267
Perdix perdix, 412
Peromyscus maniculatus, 314, 320, 412
Peromyscus maniculatus gracilis and P. leucopus nove-
boracensis, Nesting and food-storage behavior of, 239
Petrochelidon pyrrhonota, 373
Phalacrocorax carbo, 175
Phalaris arundinacea, 166
Phalarope, Red, in the prairie provinces and adjacent states,
Occurrences of the, 446
Phalaropus fulicarius, 188, 446
lobatus, 188
tricolor, 188
Phenacomys intermedius, 314
Phyllodoce coerulea, 355 —
Physalis heterophylla, 35
Physcia adscendens, 140
aipolia, 140
orbicularis, 140
Physconia grisea, 140
Picea glauca, 405
X engelmanii, 405
Pimlott, The Douglas, scholarship in environmental studies,
338
Pinguicula villosa, 264, 355
Pinicola enucleator, 436
Pinus banksiana, 282
flexilis, 405
strobus, 34
Plagiothecium denticulatum, 140
Plantago canescens, 184 |
Platydicta subtile, 140
INDEX TO VOLUME 93
473
Plethodon cinereus cinereus, 194
Pluvialis dominica, 188
squatarola, 188
Poa ammophila, 163
glauca, 184
Polygala polygama, 35
verticillata, 35
Polygonatum biflorum, 36
Polygonum amphibium, 389
amphibium var. stipulaceum, 167
lapathifolium var. lapathifolium, 378
lapathifolium var. prostratum, 379
lapathifolium var. salicifolium, 378
pensylvanicum var. durum, 379
pensylvanicum vat. laevigatum, 378
pensylvanicum var. pensylvanicum, 378
persicaria var persicaria, 379
persicaria var. ruderale, 379
scabrum, 378
Polypodiaceae, in Alberta, Taxonomy, distribution, and
ecology of the cliff-brake ferns (Pel/aea), 288
Polystichum munitum, 128
Pomoxis nigromaculatus, 232
Pontederia cordata, 389
Population densities of the amphibians of northeastern
Alberta, Habitat utilization and, 144
Populations of swallows in Canada, Man’s influence on
potential nesting sites and, 371
Populus and Salix seedfall, A trap to measure, 77
Populus balsamifera, 77
grandidentata, 34
tremuloides, 34, 77, 282
Porcupines in montane forests of southwestern Alberta,
Winter feeding by, 405
Porella platyphylla, 140
Potamogeton alpinus ssp. tenuifolius, 262
amplifolius, 388
berchtoldii, 389
epthydrus, 389
gramineus, 389
natans, 389
oakesianus, 389
obtusifolius, 388
pectinatus, 389
perfoliatus ssp. Richardsonii, 262
Richardsonii, 262, 389
robbinsii, 389
spirillus, 389
zosteriformis, 309
Potentilla elegans, 264
fruticosa, 184
hookeriana, 184
Hookeriana ssp. Hookeriana, 264
hyparctica, 264
nivea ssp. Hookeriana, 264
rubricaulis, 261
simplex, 34
Powers, K. D. Lesser Black-backed Gull, Larus fuscus, in
Labrador waters, 445
Predation on Ring-billed Gull chicks, Common Garter
Snake, 317
Predation on Robin nestlings, Common Garter Snake, 70
474
President’s message, |
Preston, J., 178
Preston, W. B., reviews by, 94, 98
Prevett, J. P., review by, 341
Prince Edward Island, 175, 325
Prionocera sp., 171
Procyon lotor, 423
Progne subis, 371
Proposals invited from field FessEe investigators, 201
Prunus americana, 36
cerasus, 34
pensylvanica, 284
serotina, 32
virginiana, 33, 284
Pseudacris triseriata maculata, 144
Pseudoleskeela catenulata, 431
Pseudotsuga menziesii, 126
menziesii var. glauca, 405
Pteridium aquilinum, 33, 128
Pteritis nodulosa, 166
Pterostichus adstrictus, 80
pensylvanicus, 80
Puffin, Horned, Nesting of, in British Columbia, 84
Puffin, Tufted, 84
Pugsley, R. W., 266
Pulsatilla patens, 184
Pylaisiella polyantha, 140
selwynii, 140
Plyodictis olivaris, 179
Pyrola secunda ssp. obtusata, 264
Pyrus coronaria, 33
Qadni, S. U., 232
Quebec, 439
Quebec, Barn Owls in, 323
Quebec, Dispersion of freshwater leeches (Hirudinoidea) to
Anticosti Island, 310
Quebec, Mosses new to Ontario and, 431
Quebec, New localities for the Northern Spring Salamander
and the Four-toed Salamander in southwestern, 193
Quercus alba, 32
bicolor, 35
macrocarpa, 34
muehlenbergii, 34
rubra, 32
velutina, 32
Quinlan, R. W. Snowy Egret in the Northwest Territories,
329
Raccoon, 423
Rana sylvatica, 144
Range extensions in eastern Canada,
niscapa at Mer Bleue and, 391
Range extensions of vascular plants in northern Yukon Ter-
ritory and northwestern District of Mackenzie, 259
Rangifer tarandus caribou, 71
t. granti, 155
Ranunculus abortivus, 167
Eschscholtzii, 263
hispidus, 36
reptans, 389
Recurvirostra americana, 68, 188
Redhorse, Black, Moxostoma duquesnei, from Ontario,
Recent collections of the, 63
Utricularia gemi-
THE-CANADIAN FIELD-NATURALIST
Vol. 93
Redhorse, Golden, 63
Shorthead, 64
Renouf, D., review by, 342
Reproductive biology of the Big Brown Bat (Eptesicus
fuscus) in Alberta, 48
Request for information — shore bird color-marking, 338
Request for participants — International Shorebird Surveys
IGA), 33s
Response of wintering Moose to mechanical habitat re-
habilitation in Alaska, 191
Responses, Demographic and dietary, of Red-tailed Hawks
during a Snowshoe Hare fluctuation, 16
Rhizomnium glabrescens, 129
Rhus typhina, 34
Rhytidiadelphus triquetrus, 129
Ribes triste, 263
Riley, J. L. Some new and interesting vascular plant
records from northern Ontario, 355
Ring counts in Salix arctica from northern Ellesmere
Island, 81
Ringius, G.S. Status of Eastern White Cedar,
occidentalis, in western Nova Scotia, 326
Riotte, J. C. E., review by, 203
Riparia riparia, 373
Ritchie, J. C., 458
Robarts, R. D., review by, 459
Roberts, W. and V. Lewin. Habitat utilization and popu-
lation densities of the amphibians of northeastern
Alberta, 144
Robin, American, 423
Robin, Common Garter Snake aredanon on nestlings, 70
Robitaille, G., reviews by, 217, 218
Roby, D: D., 155
Roosting, Communal, of Song Sparrows under snowbank,
B25
Rosa acicularis, 184
gymnocarpa, 128
spp., 284
Rowe, J. S., Is biology unknown? (letter), 330
Rubus acaulis, 264
arcticus ssp. acaulis, 264
strigosus, 284
Sagittaria graminea, 388
sp., 309
Salamander, Blue-spotted, 194
Red-backed, 194
Spotted, 194
Northern Dusky, 193
Northern Two-lined, 194
Salamander, Two-lined, Eurycea bislineata, in Labrador,
178
Salamander, Four-toed, in southwestern Quebec, New
localities for the Northern Spring Salamander and the,
193
Salamander, Northern Spring, and the Four-toed Sala-
mander in southwestern Quebec, New localities for the,
193
Salix alaxensis, 78
alaxensis var. longistylis, 167
arbusculoides, 79
Barrattiana, 167
bebbiana, 79
Thuja
979:
interior, 78
lasiandra, 79
longistylis, 167
monticola, 78
novae-anglicae, 78
padophylla, 167
planifolia, 79
spp., 79, 282
Salix arctica from northern Ellesmere Island, Ring counts
in, 81
Salix seedfall, A trap to measure Populus and, 77
Sanguisorba sitchensis, 168
stipulata, sensu, 168
Sarcophaga citellivora, 366
Saskatchewan, Colonial-nesting Herring Gulls and Com-
mon Terns in northeastern, 132
Saskatchewan, Summer, autumn, and winter diets of Elk
in, 282
Savile, D. B. O. Ring counts in Salix arctica from northern
Ellesmere Island, 81
Saxifraga caespitosa, 263
hieracifolia, 263
oppositifolia, 184
tricuspidata, 184
Scapania nemorosa, 129
Schizachne purpurascens, 166
Schmidt, D. F., 195
Scholarship, The Douglas Pimlott, in environmental studies,
338
Schowalter, D. B. and J. R. Gunson. Reproductive biology
of the Big Brown Bat (Eptesicus fuscus) in Alberta,
48
Schowalter, D. B., J. R. Gunson, and L. D. Harder. Life
history characteristics of Little Brown Bats (Myotis
lucifugus) in Alberta, 243
Schwert, D. P. and K. W. Dance. Earthworm cocoons as a
drift component in a southern Ontario stream, 180
Scirpus subterminalis, 389
Scotter, G. W. and W. J. Cody. Interesting vascular plants
from southeastern Yukon Territory, 163
Scutellaria galericulata var. pubescens, 168
Sealy, S.G., 84
Sedum rosea, 184
Selaginella selaginoides, 166
Selasphorus rufus, 450
Selkirk Mountains, 71
Senecio atropurpureus ssp. tomentosus, 265
Kjellmannii, 265
Senner, S. E. and P. G. Mickelson. Fall foods of Common
Snipe on the Copper River Delta, Alaska, 171
Sexton, D. A. and M. M. Gillespie. Effects of fire on the
location of a Sharp-tailed Grouse arena, 74
Sharp-tailed Grouse arena, Effects of fire on the location
of a, 74
Shiner, Emerald, 266
Shorebird color-marking, Request for information, 338
Shorebird Surveys, International 1979, Request for par-
ticipants, 338
Shorebirds before and since DDT, Eggshell thickness in
American, 187
Shrew, Long-tailed, 320
INDEX TO VOLUME 93 475
Masked, 314, 320
Short-tailed, 320
Smokey, 320
Water, 320
Shrew, Long-tailed, (Sorex dispar) in New Brunswick,
First record of the, 195
Sigman, M. Response of wintering Moose to mechanical
habitat rehabilitation in Alaska, 191
Silene acaulis, 184
acaulis ssp. subacaulescens, 263
menziesii, 167
Sims, R. A., 139
Skink, Five-lined, Ewmesces fasciatus, in Ontario, Eastern
limit of the, 321
Skunk, Striped, in Alberta, Winter denning of, 252
Smartweeds in Ontario, Distribution and habitats of four
annual, 378
Smelt, Rainbow, (Osmerus mordax) in Lake Simcoe,
Ontario, Food and feeding of the, 266
Smilacina racemosa, 184
trifolia, 167
Smith, L. C. Article or note? 337
Smith, L. C. Editor’s report for 1978, 201
Smith, L. C. Our responsibilities as field-naturalists and
biologists (editorial), 113 ;
Smith, L. C. Special appreciation expressed to R. Emerson
Whiting, 87
Smith, L. C. The Canadian Field-Naturalist — the status
quo or a new direction? (editorial), 10
Smith, L. C. To a bigot (editorial), 231
Smith, W. T., 155
Snake, Black Rat, 28, 322
Brown, 399
Butler’s Garter, 399
Common Garter, 70, 317
Eastern Fox, 399
Eastern Garter, 399
Northern Ribbon, 322
Snake, Common Garter, predation on Robin nestlings, 70
Snakes at Amhertsburg, Ontario, Movements of sympatric
species of, 399
Snipe, Common, Fall foods of, on the Copper River Delta,
Alaska, 171
Snowshoe Hare fluctuation, Demographic and dietary
responses of Red-tailed Hawks during, 16
Solidago caesia, 33
canadensis, 35
canadensis var. salebrosa, 169
juncea, 35
multiradiata, 184
nemoralis, 35
Sonchus asper, 284
Song pattern of the Cypress Hills population of White-
crowned Sparrows, 272
Sorex arcticus, 315
cinereus, 314, 320
dispar, 320
fumeus, 320
palustris, 315, 320
Sorex dispar, First record of the Long-tailed Shrew, in
New Brunswick, 195
476
Sparganium angustifolium, 388
fluctuans, 389
minimum, 262
Sparrows, Song, under snowbank, Communal roosting of,
395
Sparrows, White-crowned, Song pattern of the Cypress
Hills population of, 272
Spermophilus franklinii, 20
richardsonii, 17, 363, 412
Sphaeroderus nitidicollis, 80
Sphenopholis intermedia, 163
Spiranthes Romanzoffiana, 263
Squirrel, Franklin’s Ground, 20
Northern Flying, 320
Red, 320
Richardson’s Ground, 17, 19, 412
Squirrels, Richardson’s Ground, Yearly variations in the
population dynamics of, 363
Staniforth, R. J. and P. B. Cavers. Distribution and habitats
of four annual smartweeds in Ontario, 378
Stelfox, H. A. and G. J. Brewster. Colonial-nesting Herring
Gulls and Common Terns in northeastern Saskatche-
wan, 132
Stelgidopteryx ruficollis, 372
Stellula calliope, 449
Stepney, P. H.R. Brewer’s Blackbird breeding in the
Northwest Territories, 76
Sterna hirundo, 132
Sternotherus odoratus, 322
Stilts, Black-necked, nesting near Edmonton, Alberta, 68
Stinkpot, 322
Stocek, R. F. Decline of summering Bald Eagles in central
New Brunswick, 443
Storeria dekayi, 399
Storey, A., 303
Sullivan, M.G. Blue Grouse brood hen — Black Bear
confrontation, 200
Sullivan, T. P. Virgin Douglas Fir forest on Saturna Island,
British Columbia, 126
Swallow, Bank, 373
Barn, 374
Cliff, 373
Rough-winged, 372
dinees 372
Violet-green, 372
Swallows in Canada, Man’s influence on potential nesting
sites and populations of, 371
Swallows, Tree, Influence of weather on aggression in, 437
Swan, Mute, 433
Trumpeter, 433
Swans wintering on Vancouver Island, 1977-1978, 433
Swertia caroliniensis, 36
Systematic and evolutionary biology, Second International
congress of, 87
Tachycineta thalassina, 372
Tadlock, C. G. and H. G. Klein. Nesting and food-storage
behavior of Peromyscus maniculatus gracilis and P.
leucopus noveboracensis, 239
Tamias striatus, 315, 320
Tamiasciurus hudsonicus, 320
Taraxacum officinale, 184
Phymatocarpum, 265
THE CANADIAN FIELD-NATURALIST
Vol. 93
Taxidea taxus, 365
Taxonomy, distribution, and ecology of the cliff-brake ferns
(Pellaea: Polypodiaceae) in Alberta, 288
Taylor, P. Interspecific vocal mimicry by Pine Grosbeaks,
436
Teal, Blue-winged, 316
Cinnamon, 316
Teal, Cinnamon X Blue-winged, from southern Alberta,
Probable hybrids of, 316
Tendipes tentans, 267
Terasmae, J. and N. C. Weeks. Natural fires as an index
of paleoclimate, 116
Terns, Black, Nesting biology and development of young in
Ontario, 276
Terns, Common, in northeastern Saskatchewan, Colonial-
nesting Herring Gulls and, 132
Thalictrum sparsiflorum var. richardsonii, 167
Thamnophis butleri, 399
Sauritus septentrionalis, 322
sirtalis, 70, 317
sirtalis sirtalis, 399
Thaspium barbinode, 36
Thaspium trifoliatum (Meadow-parsnip) in Canada, 306
Thompson, I. D., review by, 341
Thomson, S. C., review by, 101
Thormin, T. W., 68
Thuja occidentalis, in western Nova Scotia, Status of
Eastern White Cedar, 326
Thuja plicata, 126
Tilia americana, 34
To a bigot (editorial), 231
To know ourselves, 89
Toad, Canadian, 144
Todd, A. W., 16
Townsendia hookeri, 184
Trans-Alaska Pipeline, Caribou distribution and group
composition associated with construction of the, 155
Trientalis latifolia, 129
Triticum aestivum, 285
Tsuga heterophylla, 126
Turdus migratorius, 70, 423
Tyto alba, 323
Ulmus americana, 34
Ulmus americana, Epiphytes on White Elm near Thunder
Bay, Ontario, 139
Urquhart, F. A. and N. R. Urquhart. Breeding areas and
overnight roosting locations in the northern range of the
Monarch Butterfly (Danaus plexippus plexippus) with
a summary of associated migration routes, 41
Urquhart, N. R., 41
Ursus americanus, 200, 423
Urtica gracilis, 167
Ussher, R. D. and F. R. Cook. Eastern limit of the Five-
lined skink, Eumesces fasciatus, 1n Ontario, 321
Utricularia cornuta, 388
gibba, 389
intermedia, 388
purpurea, 388
resupinata, 388
vulgaris, 264, 389
Utricularia geminiscapa at Mer Bleue and range extensions
in eastern Canada, 391
IVE
Vaccinium angustifolium forma angustifolium, 416
angustifolium forma leucocarpum, 416
angustifolium forma nigrum, 416
pallidum, 36
vacillans, 36
vitis-idaea, 184
Vahlodea atropurpurea, 355
Vallisneria americana, 389
Vancouver Island, 1977-1978, Swans wintering on, 433
Vander Kloet, S. P., 415
Vanellus chilensis, 189
Vascular plant records from northern Ontario, Some new
and interesting, 355
Vascular plants from southeastern Yukon Territory, Inter-
esting, 163
Vascular plants in northern Yukon Territory and north-
western District of Mackenzie, Range extensions of,
259
Viburnum acerifolium, 33
edule, 184
trilobum, 284
Vicia americana, 163, 284
Viola langsdorffii, 168
pedata var. lineariloba, 36
rugulosa, 163
Vole, Heather, 314
Meadow, 18, 320, 412
Red-backed, 314, 320
Rock, 314, 320
Vole, Rock (Microtus chrotorrhinus) as a Transition zone
species, The, 319
Vulpes vulpes, 423
Wapiti, 282
Weasel, Long-tailed, 365, 412
Weasel, Long-tailed, and Snowy Owl, Interaction between
a, 67
INDEX TO VOLUME 93
477
Weather, Influence of, on aggression in Tree Swallows, 437
Webber, G. A., 176
Weber, W. C., reviews by, 91, 203
Weeks, N. C., 116
Weseloh, D. V., 68
Weseloh, D. V. and L. M. Weseloh. Probable hybrids of
Cinnamon X Blue-winged Teal from southern Alberta,
316
Weseloh, D. V., review by, 96
Weseloh, L. M., 68, 316
Whale management, IUCN views on, 88
White, E. R., 82
Whiting, R. Emerson, Special appreciation expressed to, 87
Whitten, K R., 155
Wildlife film festival, second annual international, 90
Winder, G., 28
Wolf, Gray, skulls found in Alberta, Largest, 308
Woodchuck, Aquatic feeding by a, 309
Wright, D. G., 199
Xanthochroism in the Evening Grosbeak, 66
Xanthoria polycarpa, 140
Young, J. W.S. review by, 348
Yukon Territory and northwestern District of Mackenzie,
Range extensions of vascular plants in northern, 259
Yukon Territory, Flowering plant phenology at Sheep
Mountain, southwest, 183
Yukon Territory, Interesting vascular plants from south-
eastern, 163
Zammuto, R. M., review by, 461
Zapus hudsonius, 314, 320
Zasada, J.C. and R. Densmore. A trap to measure Populus
and Salix seedfall, 77
Zizania aquatica, 389
Zonotrichia leucophrys oriantha, 272
Zygadenus elegans, 184
Index to Book Reviews
Botany
Arditti, J. (ed.). Orchid biology: reviews and perspectives,
1., 104
Argus, G. W. and D. J. White. The rare vascular plants
of Ontario/Les plantes vasculaires rares de Ontario,
100
Bassett, I. J. An atlas of airborne pollen grains and
common spores of Canada, 457
Brodie, H. J. Fungi: delight of curiosity, 101
Brunton, D. F. Flora of Alberta: a checklist, 213
Cody, W. J. Ferns of the Ottawa district, 345
Dorn, R. D. Manual of vascular plants of Wyoming, 211
Good, R.E., D. F. Wigham, R. L. Simpson, and C. G.
Jackson, Jr. (eds.). Freshwater wetlands: ecological
processes and management potential, 213
Harrington. H. D. How to identify grasses and grasslike
plants (sedges and rushes), 212
Markgraf, V.and H. L. D’Antoni. Pollen flora of Argentina:
modern spore and pollen types of Pteridophyta, Gymn-
nospermae, and Angiospermae, 102
McGrath, J. W. Dyes from lichens and plants: a Canadian
dyer’s guide, 346
Muenscher, W. C. and M. A. Rice. Garden spice and wild
pot-herbs: an American herbal, 212
Nilsson, S. Atlas of airborne pollen grains and spores in
northern Europe, 103
Shetler, S. G. and L. E. Skog (eds.). A provisional checklist
of species for Flora North America (revised), 212
Smith, J. P., Jr. Vascular plant families, 457
Szezawinski, A. F. and N. J. Turner. Edible garden weeds
of Canada, 344
Zander, R.H. Floristics and environmental planning in
western New York and adjacent Ontario: distribution of
legally protected plants and plant sanctuaries, 102
Environment
Bow Valley Naturalists. Vermillion Lakes, Banff National
Park: an introductory study, 347
Button, J. The Shetland way of oil: reactions of a small
community to big business, 105
Colinvaux, P. Why big fierce animals are rare: an ecologist’s
perspective, 108
478
Dansereau, P. EZAIM: écologie de la zone de l’aéroport
international de Montréal — le cadre d’une recherche
écologique interdisciplinaire, 106
Girard, C. M. and M.C. Applications de la télédétection
a l'étude de la biosphere, 217
Jurant, M., J. L. Bélair, V. Gerardin, and J. P. Ducrue.
L’inventaire du capital nature: méthode de classification
et de cartographie écologique du territoire (3 éme
approximation), 216
Kinkead, E. Wildness is all around us:
urban naturalist, 217
Long, D. R. (ed.). Outdoors Canada: a unique and practical
guide to our wilderness and wildlife, 107
McMillan, V. and B. Nature quizzes for Canadians, 104
Pruitt, W. O. Boreal ecology, 215
Ramel, C. (ed.). Chlorinated phenoxy acids and their
dioxins, 460
Russell, D. A. A vanished world: the dinosaurs of western
Canada, 458
Shelford, V. E. The ecology of North America, 460
Statistics Canada. Human activity and the environment, 347
Wall, G. and C. Wright. The environmental impact of out-
door recreation, 215
Werger, M.J..A (ed.). Biogeography and ecology of
southern Africa, 459
notes of an
Zoology
Beebe, F. L. Hawks, falcons and falconry, 205
Blokpoel, H. Bird hazards to aircraft: problem and pre-
vention of bird/ aircraft collision, 96
Brownie, C., D. R. Anderson, K. P. Burnham, and D.S.
Robson. Statistical inference from band recovery data:
a handbook, 455
Bruemmer, F. Life of the Harp Seal, 342
Cayouette, R. Nichoirs d’oiseaux, 210
Clark, J. L. The great arc of the wild sheep, 339
Comstock, A. B. Ways of the six-footed, 94
deSchauensee, R. M. and W. H. Phelps, Jr. A guide to the
birds of Venezuela, 340
Eriksson, P. S. The bird finder’s 3-year note book, 99
Erskine, A. J. Birds in boreal Canada, 207
Ferguson, D. C. The moths of America north of Mexico,
including Greenland. Fascicle 22.2, Noctuidae (in part):
Lymantriidae, 203
Goodwin, D. Birds of Man’s world, 454
Goodwin, D. Crows of the world, 342 -
Grainger, D. C. Animals in peril: a guide to the endangered
species of Canada and the United States, 339
THE CANADIAN FIELD-NATURALIST
Vol. 93
Grassé, P. P. Précis de zoologie: vertébrés. 3 — repro-
duction, biologie, évolution et systématique. Oiseaux et
mammiféres, 94
Harington, C. R. Quaternary vertebrate faunas of Canada
and Alaska and their suggested chronological sequence,
453
Kelley, A. H. Birds of southeastern Michigan and south-
western Ontario, 95
Mackenzie, J. P.S. Birds in peril: a guide to the en-
dangered birds of Canada and the United States, 210
Mark, D. M. Where to find birds in British Columbia, 203
Merritt, R. W. and K. W. Cummins (eds.). An introduction
to the aquatic insects of North America, 209
Nelson, B. The Gannet, 456
Nelson, J. S. Fishes of the world, 206
Ogilvie, M. A. Wild geese, 341
Pelton, M.R., J. W. Lentfer, and G.E. Folk (eds.).
Bears: their biology and management, 97
Perry, R. Watching sea birds, 95
Peterson, R.L. North American Moose, 341
Pettingill, O.S., Jr. A guide to bird finding east of the
Mississippi, 91
Raveling, D. G. and H. G. Lumsden. Nesting ecology of
Canada Geese in the Hudson Bay Lowlands of Ontario:
evolution and population regulation, 207
Reynolds, J. W. The earthworms (Lumbricidae and Spar-
ganophilidae) of Ontario, 98
Schaller, G. B. Mountain monarchs: wild sheep and goats
of the Himalaya, 93
Schorger, A. W. The Passenger Pigeon: its natural history
and extinction, 98
Squires, W. A. The birds of New Brunswick, 92
Tanner, J. T. Guide to the study of animal populations, 453
Vaillancourt, J. Lexique anglais-frangais: termes techniques
a lusage des biologistes, 208
Miscellaneous
Batschelet, E. Introduction to mathematics for life scientists,
349
Blaker, A. A. Field photography: beginning and advanced
techniques, 218
Corrivault, G.-W. and P. Morisset. Le Naturaliste Canadien,
Index, 462
Inglis, A. Northern vagabond: the life and career of
J. B. Tyrrell, 109
Ruffner, J. A. and F. E. Bair (eds.). Weather almanac, 348
Sterling, K. B. Last of the naturalists: the career of C. Hart
Merriam, 461
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TABLE OF CONTENTS (concluded)
Book Reviews
Zoology: Guide to the study of animal populations — Quaternary vertebrate faunas of Canada 453
and Alaska and their suggested chronological sequence — Birds of Man’s world —
Statistical inference from band recovery data: a handbook — The Gannet
Botany: Vascular plant families — An atlas of airborne pollen grains and common spores of Canada 457
Environment: A vanished world: the dinosaurs of western Canada — Biogeography and ecology of 458
southern Africa — The ecology of North America — Chlorinated phenoxy acids and their
dioxins
Miscellaneous: Last of the naturalists: the career of C. Hart Merriam — Le Naturaliste 461
Canadien Index
New Titles 463
Index to Volume 93 Compiled by W. HARVEY BECK 465
Mailing date of previous issue 26 June 1979
Erratum
Canadian Field- Naturalist 93(3): 232-238; 1979.
Seasonal growth, food, and feeding habits of young-of-the-year Black Crappie in the Ottawa River by John
Mark Hanson and S. U. Qadri.
The log,, values for total length given on the X-axis of Figure 4 are incorrect; they should be 1.1, 1.2, 1.3, 1.4, 1.5,
RGR plevenlese01-9 > and)2-0!
1979 Council — The Ottawa Field-Naturalists’ Club
President: Roger Taylor Elisabeth Beaubien Fran Goodspeed
Charlie Beddoe Peter Hall
Vice-President: Courtney Gilliatt Ron Bedford Hue MacKenzie
¢ Frank Bell Frank Pope
Treasurer: Barry Henson Bill Cody eee eaee
Recording Secretary: Diana Laubitz Jane Diceman Ken Strang
; . Ellaine Dickson Ken Taylor
Corresponding Secretary: Valerie Hume oneuneKson ee
Past President: Roger Foxall Marc Forget
Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists’ Club,
Box 3264, Postal Station C, Ottawa, Canada K1Y 4J5. For information on Club activities telephone (613) 722-3050.
THE CANADIAN FIELD-NATURALIST Volume 93, Number 4 1979
i
Articles
Some new and interesting vascular plant records from northern Ontario J. LL REE
Yearly variations in the population dynamics of Richardson’s
Ground Squirrels GAIL R. MICHENER
Man’s influence on potential nesting sites and populations of swallows
in Canada A. J. ERSKINE
Distribution and habitats of four annual smartweeds in Ontario
RICHARD J. STANIFORTH and PAUL B. CAVERS
Apparent differences in aquatic macrophyte floras of eight lakes in Muskoka
District, Ontario from 1953 to 1977 G. E. MILLER and H. M. DALE
Utricularia geminiscapa at Mer Bleue and range extensions in
eastern Canada ERICH HABER
Movements of sympatric species of snakes at Amherstburg, Ontario
B. FREEDMAN and P. M. CATLING
Winter feeding by Porcupines in montane forests of southwestern
Alberta LAWRENCE D. HARDER
Interactions between Snowy and Short-eared Owls in winter
M. Ross LEIN and PETER C. BOXALL
The Biological Flora of Canada
1. Vaccinium angustifolium Ait., Sweet Lowbush Blueberry
IVAN V. HALL, LEwis E. AALDERS, NANCY L. NICKERSON, and SAM P. VANDER KLOET
Notes
Mosses new to Ontario and Quebec ROBERT R. IRELAND and GILDA BELLOLIO-ITRUCCO
Swans wintering on Vancouver Island, 1977-1978 RICHARD W. MCKELVEY
Interspecific vocal mimicry by Pine Grosbeaks PETER TAYLOR
Influence of weather on aggression in Tree Swallows REID N. HARRIS
Abandoned windmill used as a nesting site by Great Blue Herons JEAN-LUC DESGRANGES
Unusually late pregnancy of a Muskrat in southeastern New Brunswick G. R. PARKER
Lynx movements and habitat use in Montana
GARY M. KOEHLER, MAURICE G. PeRNaceR. and HOWARD S. HASH
Decline of summering Bald Eagles in central New Brunswick Re PB; SiOGEK
Lesser Black-backed Gull, Larus fuscus, in Labrador waters KEVIN D. POWERS
Occurrence of the Red Phalarope in the Canadian prairie provinces and
adjacent states J. PAUL GOOSSEN and DANIEL G. BUSBY
Nesting of the Calliope Hummingbird in Kananaskis Provincial Park, Alberta
DANIEL F. BRUNTON, SIDNEY ANDREWS, and DAVID G. PATON
News and Comment
391
399
405
411
415
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433
436
437
439
440
44]
443
445
446
449
452
concluded on inside back cover
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